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Silurian paulinitid polychaetes from Gotland CLAES F. BERGMAN

Bergman, Claes F. 1989 Il 30: Silurian paulinitid polyehaetes from Gotland. Fossils and Strata, No. 25, pp. 1-1 28. Oslo. ISSN 0300-9491 . ISBN 82-00-37424-6.

Silurian paulinitids eonstitute a large and diverse group of jawed polyehaetes whieh flourished in the tropieal epicontinental sea in the Gotland area. They are represented by five genera: Gotlandites, Hindenites, Lanceolatites, Kettnerites [with the subgenus K (Aeolus)], and Langeites. Twenty species and nine subspeeies and varieties from latest Llandovery to Late Ludlow are identified, based on apparatuses reconstrueted by utilizing isolated elements (seoleeodonts). A biologieaI spe eies eoneept is employed, ineorporating taxa based on jaw elements as weU as on apparatuses. The loeal temporai and geographie oeeurrenees of the paulinitids on Gotland are established. Some lineages evolve very slowly throughout the sequenee, and one group forms a complex of shott lineages. A third eategory of taxa is found only in speeifie environments. It is eoncluded that at least some spe eies are intraeontinentaIly distributed. The study is based on severai tens of thousands of jaws from more than 700 samples from 342 loealities. DPolychaetes, eunicids, paulinitids, jaw apparatuses, scolecodonts, taxonomy, phylogeny, ontogeny, palaeogeographical distrilmtion, Silurian, Gotland, Sweden, N5654 N5800 E1921 E1758.

Claes F. Bergman, Department of Historical Geology and Palaeontology, University of Lund, Sijlvegatan 13, S-223 62 Lund, Sweden; 1988 02 16.

Contents Introduetion ................ . . ................. 3 Historical review ................................. 4

Accounts of fossil annelidjaws fromSweden ....... 6 Geology of Gotland . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 6

Stratigraphical and geographical distribution of paulinitids on Gotland ...................... 7

Ecological notes ............................... 25 Comparison with paulinitid faunas from other areas . 26 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . .. 27

Illustration of speeimens . . . . . . . . . . . . . . . . . . . . . .. 28 Terminology ................................ 29

Structure, composition, and preservation of polychaetejaws .............................. 31

Molting ofjaws and ontogeny .............. . ..... 33 General taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 34

Variability ofjaws and apparatuses ............... 34 Morphological features of taxonomical

importance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 35 Scolecodonts versus apparatuses ................ 35

Classification .................................. 39 Phylum Annelida Lamarek 1809 ................ 39 Class Polychaeta Grube 1850 ................... 39 Order Eunicida Dales 1963 . . . . . . . . . . . . . . . . . . . .. 39 Superfamily Eunicea Grube 1852 . . . . . . . . . . . . . . .. 39 Family Paulinitidae Lange 1947 ................. 39 Genus Gotlandites Bergman 1987 ................ 39

Gotlandites slitensis Bergman 1987 . . . . . . . . . . . . .. 40 Genus Hindenites Bergman 1987 ................ 43

Hindenites angustus (Hinde 1882) .............. 43

Hindenites gladiatus (Kielan:J aworowska 1966) ... 45 Hindenites naerensis Bergman 1987 . . . . . . . . . . . .. 47

Genus Kettnerites Zebera 1935 .................. 49 Kettnerites (K) abraham Bergman 1987 . . . . . . . .. 49 Kettnerites (K) abraham abraham Bergman 1987 .. 51 Kettnerites (K) abraham isaac Bergman 1987 . . . .. 53 Kettnerites (K) jacobi Bergman 1987 ........... 54 Kettnerites (K) bankvaetensisBergman 1987 ..... 57 Kettnerites (K) burgensis (Martinsson 1960) ..... 62 Kettnerites (K) huberti Bergman 1987 .......... 65 Kettnerites (K) martinssonii Bergman 1987 ...... 70 Kettnerites (K) polonensis (Kielan:Jaworowska

1966) ....................... . . . . . . . . . .. 76 Kettnerites (K) polonensisvar. sjaustre .......... 82 Kettnerites (K) sp. A ........................ 84 Kettnerites (K) versabilis Bergman 1987 . . . . . . . .. 85 Kettnerites (Aeolus) Bergman 1987 . . . . . . . . . . . .. 87 Kettnerites (Aeolus) sisyphi Bergman 1987 ....... 88 Kettnerites (Aeolus) sisyphi sisyphi Bergman 1987 .. 89 Kettnerites (Aeolus) sisyphi var. valle ............ 92 Kettnerites (Aeolus) sisyphi klasaardensis Bergman

1987 . .. . . . . . . . . . . . . . . . . . . . . . .. . . . .. . . .. 94 Kettnerites (Aeolus) microdentatus Bergman 1987 .. 99 Kettnerites (Aeolus) siaelsoeensis Bergman 1987 .. 102 Kettnerites (Aeolus) Jjaelensis Bergman 1987 . . . .. 104

Genus Lanceolatites Bergman 1987 ............. 106 Lanceolatites gracilis Bergman 1987 ........... 106 Lanceolatites gracilis var. visby . . . . . . . . . . . . . . .. 11 O Lanceolatites sp. A ................ . . . ...... I I I

Genus Langeites Kielan:Jaworowska 1966 ........ 112 Langeites glaber Kielan:J aworowska 1966 ....... 112

Referenees .................................. 113 Appendix: Localities .......................... 119

Introduction Polychaete jaw apparatuses of quite different types have

developed in the widespread Recent errant taxa of the

Phyllodoeida and the Eunicida. In the Phyllodocida (in

cluding, e.g., Nereidae, Nephtyidae, and Glyceridae) the

proboscis is axial. In the Eunicida (e.g. Eunicidae, Lumbri

nereidae, Onuphidae, Dorvilleidae, and the extinct Pauli

nitidae) the proboseis is in the ventrai position and armed

with a more complexjaw apparatus than in the Phyllodoc

ida.

It has been my ambition to employ a neontological spe

eies concept in the present work and to introduce a nomen

clature which follows the ICZN rules. Thus, the two paralleI

taxonomical systems currently employed for fossil poly

chaetes, the form-taxonomic (usually based on isolated

jaws) and what is known as the natural taxonomic (usually

based on jaw apparatuses) systems are here merged into

one.

The excellent conditions of preservation for the organic

polychaete jaws in the Silurian sediments of Gotland

greatly facilitated the present reconstruction work. Fur

ther, the Silurian marine, tropical, shallow shelf seas have

offered a rich variation of habitats within different types of

environments e.g. reefs, lagoons and open marine soft and

hard bottoms. Thus, with dense sampling, it has been

possible to study the evolutionary trends of populations as

well as the preference of the polychaetes for different

environments (i.e. the jaws have been found in a particular

sediment).

The form taxonomic system has plagued studies of fossil

jawed annelids ever since Hinde published his papers on

annelidjaws towards the end of the 19th century. Based on

a statement on Recent jaws by Claparede (1870, p. 24),

Hinde assumed that the fossil jaws would also be useless in

the taxonomical work and was conseious of the fact that his

taxonomy was tentative (Hinde 1880, pp. 369-370). On the

other hand, Hinde (1879) established the genus Arabellites

to include severai different jaws by analogy with the jaw

apparatus of the Recent genus Arabella.

The state of affairs is similar to that of conodont taxon

omy until the midd le-late part of 1960's. Today most con 0-

dontologists have accepted the biological taxonomical con

cept, initiated by the works of Bergstrom & Sweet (1966),

Webers (l966) ,]eppsson (1969 and 1974) , and von Bitter

(1972) , as the on ly practical solution.

Finds of complete polychaete jaw apparatuses are valu-"

able, particularly for reconstructions, which include the

identification of the type, num ber and arrangement of the

elements. In spite of the large num ber of polychaete jaw

4 Claes F. Bergman

apparatuses described, the polychaete taxonomy is in a bad

state. One explanation may be that the scolecodonts are

of ten more or less deformed, making identification of sin

gle elements and apparatuses difficult.

Among scolecodonts, a large number of names based on

form-taxonomy of different jaw elements must be revised,

and the apparatus-based names must be included in the

revision. Severai students of scolecodonts would probably

agree on the desirability of this (e.g. Szaniawski & Wrona

1973) but only a few have been more or less successful in

their attempts to deal with the problem (e.g. Mannil & Zaslawskaya 1985a). The attempt by Kozur (1970) to com

bine the two taxonomical systems into one was premature.

Still, it is possible to identify species on different single jaw

elements, at least among the labidognatha (e.g. Kielan

Jaworowska 1966, pp. 40-42). However, scolecodontolo

gists have not been convinced of the advantage of a merge

of the two taxonomical systems.

The Gordian knot of the polychaete parataxonomy

seems to be the fact that it might not be possible to identify

all dispersed elements forming an apparatus. This obstacle

may perhaps never be overcome but it must be much more

important to distinguish the different taxa on the specific

level even if only one element is identifiable in an appara

tus, than to 'identify' broad form categories of no use for

any other studies. Thus, the solution is simply to cut the

knot and, at present, forget the identification of for exam

ple small anterior elements, since it is not of primary inter

est to identify all elements in an apparatus at the specific

leve!.

A taxonomy based on populations is the platform for

further work. The corresponding nomenclature includes

names from both taxonomical systems. A sound species

concept will without doubt facilitate stratigraphical, taxo

nomical, ecological, geographical and phylogenetical stud

ies and make the polychaete jaws useful as tools for environ

mental interpretations and as index fossils.

An earlier version of the present monograph was printed

and distributed in a limited edition (270 copies) in August,

1987, to meet the requirements for a Swedish Ph.D. thesis

examination. Unfortunately, that version was not provided

with a statement disclaiming the validity of the new system

atic names (ICZN, Article 8b). Furthermore, it has been

obtainable on request from the University of Lund. Conse

quently, the new taxonomic names must be treated as

having been made available in 1987. Due to the limited

availability of the thesis edition, full taxonomic information

regarding these 1987 names is included in the present

published version of the monograph.

Acknowledgements. - Much of my work has been carried out at the Department ofHistorical Geology and Palaeontology, University of Lund. During this time Gerhard RegneIl, Anita Liifgren, and Kent Larsson (present Head of the Department) have provided me with working facilities.

Sven Laufeld suggested the topic and introduced me to the geology of Gotland. Anita Liifgren has read the various drafts of the final version of this monograph and she has generously given me a never failing support in my work from strict editorial aspects to taxonomical considerations. Her comments have greatly improved this manuscript. Enrico Serpagli, Hubert Szaniawski and Stefan

FOSSILS AND STRATA 25 (1989)

Bengtson have scrutinized the manuscript and suggested severaI most valuable improvements to it.

Grants to Anita Liifgren from the Swedish NaturaI Science Research Council (NFR) (G-GU 4746-100, 1 03, 1 04, 105, IlO) and to Kent Larsson (G-GU 3676-102, 103, 107) have financed the main part of this project. Most of the extensive field work has been carried out under the auspices of Project Ecostratigraphy paid through the NFR. Lunds GeologiskaFiiltklubband Th. NordstromsFond supported part of the field work on Gotland. The Royal Swedish Academy of Science arranged, through the Polish Academy of Science, the possibility to study the extensive paulinitid collection deposited in the Palaeozoological Institute of the Polish Academy of Science, Warsaw. Travel costs to Poland have been paid by Hans Emil Hanssons Fond. Travel to Great Britain has been funded by grants from von Beskows Fond.

Gerhard RegnelI has given me information concerning the early Swedish literature. Birger Berg has corrected my latinized taxonomical names. Kristina Lindholm translated parts of the Russian papers.

Takeshi Miyazu gave technical assistance when the SEM equipment failed to follow my intentions. Sven Stridsberg coated the SEM preparates. It would have been almost impossible to process the samples without technical assistance, paid for through grants from NFR. SeveraI of the sample residues derive from Lennart

J eppsson' s samples, most of which have been processed by him and his staff. The following persons have carried out the laboratory handling of the sam pl es, mostly involving the picking out of scolecodonts from the residues: Bjiirn Olof Gustavsson, Marie Jiinsson, Peter Mileson, Sara Nyman, Anja Rosenberg, Ewa Sall, and Cecilia Wieslander. Doris Fredholm has given me a helpful hand now and then by supervising my laboratory staff and a1so by checking my locality list. Three of my samples were processed by staff employed at the department.

The following colleagues have generously given me access to their samples: LennartJeppsson, Doris Fredholm, Kent Larsson, Sven Laufeld, Louis Liljedahl and Sven Stridsberg.

Material from the Vattenfallet section at the Swedish Museum of Natural History, Stockholm was lent by Valdar Jaanusson.

Access to the Hinde material at the British Museum (NaturaI History), London, was given by D. L. F. Sealy, that ofSnajder at the Narodny Museum, Prag, by Rudolf Prokop and that of Kielan

Jaworowska and Szaniawski in the Palaeozoological Department in Warsaw by Hubert Szaniawski.

Finally, particular thanks are due to LennartJeppsson. He initiated the study, and has put considerable time and effort into this project. Recurrent discussions with him on the geology of Gotland, apparatus reconstructions and on various zoological aspects, from vertebrates to invertebrates have been most inspiring. His ideas and advice have be en very valuable to me and his comments on the manuscript have greatly improved it.

To all these friends, colleagues, students, not forgetting the participants in the Project Ecostratigraphy and other persons who have supported and contributed to my study, I am deeply thankful.

Historical review Among the earliest published accounts of jawed annelids

are those by Eichwald (1854), Massalongo (1855), and

Pander (1856), issued more or less simultaneously without

the authors' being aware of each others' work. Both Eich

wald and Pan der described isolated annelid jaws from the

Lower Palaeozoic of the Baltic region, referring to them as

denticles of fish. Massalongo on the other hand described

impressions of annelids with the jaws preserved in place,

from Tertiary beds in Italy; thus he could identify the jaws

as being those of polychaetes. His monograph was probably

not widely circulated, and when Hinde worked with the


Lower Palaeozoic jaws decades later he was not aware of it.

Thus, Hinde (1882, p. 4) and Thorell & Lindstr6m (1885,

p. 4) give N. P. Angelin, a Swedish palaeontologist, the

credit for be ing the first to place fossil polychaete jaws in

their proper taxonomic place. They refer to a letter written

in 1864 by Angelin in which he has correctly identified

annelid jaws. Like Massalongo, the German zoologist

Ehlers (1864-68, 1868a, 1868b) described imprints of an

nelid bo dies and jaws, but the jaws were in a bad state of

preservation in Ehler's Solnhofen material. As opposed to

Massalongo and Ehlers, Grinnell (1877) worked with iso

lated elements. He based a new annelid genus on a poorly

preserved jaw fragment later regarded as a nomen dubium

(Kozur 1970, p. 44) .

Although not be ing the first to describe fossil annelid

remains, Hinde must be regarded as one of the pioneers of

scolecodont research with four contributions on Early Pal

aeozoic annelidjaws (1879, 1880, 1882, and 1896) . Hinde

was aware of the affinity of the jaws, and knew that in recent

eunicid annelids the buccal armatures are differentiated

with various elements composing an apparatus. In spite of

this, Hinde worked both with an apparatus-based and with

a morphological-based taxonomy. However, he notes

(1880, pp. 369-370) that his work is of a tentative nature.

The explanation of this treatment is found in a paper by

Claparede, one of the leading authorities on Recent anne

lids at that time, who stated (1870, p. 24) that it was

impossible to use the jaws of annelids for any taxonomic

work. Thus, the myth that it is impossible to use isolated

annelid jaws for the identification ·of 'true' species was bom

and the concept of morphological species was initiated

which has since then be en used by several later scolecodont

workers.

Over the next decades, at the beginning of the twentieth

century, very few reports on fossil wormjaws appeared. At

the beginning of the 1930's however, the time seerned right

for further studies. Still, each jaw type was treated as de

rived from a separate species. In an abstract Croneis & Scott

(1933) denominated isolated polychaete jaws 'scoleco

donts', and this term has won full acceptance and is widely

used. According to Jansonius & Craig (1971, p. 252) , Cro

neis & Scott supervised severai doctoral theses on annelid

jaws but published only the abstract of 1933 themselves. In

the same year both Eller (1933) and Stauffer (1933) made

their first contributions to scolecodont research. Eller con

tinued to publish papers on scolecodonts until 1969, with

more than 20 accounts dealing with Palaeozoic scoleco

donts. Zebera (1935) described and discussed isolated sco

lecodonts and scolecodonts in clusters from the Palaeozoic

strata of Bohemia.

Zebera's material was later revised by Snajdr (1951) who

worked with a multielement species concept and by so

doing is one of the first palaeontologists to apply a naturai

taxonomy to scolecodonts.

From Devonian strata of Parana, Brazil, Lange (1947)

recorded more or less complete polychaete jaw appara

tuses on bedding planes. His penetrating study included a

large num ber of apparatuses as well as comparative studies

on Recent material. Lange concluded that there is a large

Silurian paulinitid polychaetes from Gotland 5

variation in the morphology of the jaws in apparatuses of

Recent annelids. Furthermore, he also referred to the liter

ature on fossil jaws and singled out a number of isolated

scolecodonts resembling the ones in his material. The

identifications were based on illustrations and because of

the confused taxonomic situation of scolecodonts and the

lack of knowledge of other fossil anne lid apparatuses,

Lange (1949, pp. 48-56) noted that his synonymy list was

tentative. As he was uncertain of the relationship between

the scolecodonts described earlier and his material, he

found it safer to name a new genus based on the appara

tuses. This has had a major impact on subsequent fossil

annelid research, because the parataxonomic system evol

ved out of this work. This system included the morpholog

ically based system initiated by Hinde for dispersed jaws

(scolecodonts) on the one hand, and the new biological

system for apparatuses on the other. It has since then been

very tempting for students working with apparatuses to use

this apparatus-based taxonomy only, as there is then no

pressure to work with the confused form-taxonomic scolec

odont system. Later, Jansonius & Craig (1971) tried to sort

out the scolecodont taxonomy, i.e. they grouped the ele

ments in form-taxonomic categories. Because of the lack of

information on the isolatedjaws (e.g. the specific variation

of the elements and the composition of apparatuses, etc.) ,

they did not attempt to make a true biological taxonomy

that should also have included the apparatus-based taxa in

their work. ButJansonius & Craig (1971, p. 253) noted that

with additional information it could be anticipated that

eventually all scoJecodonts can be assigned to a genus in

accordance with the rules of ICZN.

Reconstructions of fossil polychaete jaw apparatuses

from isolated elements have been made and discussed in

severai other studies and by different students, e.g. Sylves

ter (1959) , Kielanjaworowska (1966) , Szaniawski (1968) ,

Kozur (1971) , Corradini & Olivieri (1974) , Jansonius & Craig (1974) , Szaniawski & Gazdzicki (1978) , Bergman

(1979,1980, and 1981b) , Mannil & Zaslavskaya (1985a, bl.

Their approaches to achieve a correct reconstruction rep

resent different and feasible ways based on the available

material (see the chapter 'Descriptions and reconstruc

tions of annelid jaw apparatuses') .

The idea of working with the apparatus-based taxonomy,

the natural taxonomy introduced by Lange (1947, 1949), was accepted and used by severai polychaete workers e.g.

Kozlowski (1956) , Martinsson (1960) , Kielanjaworowska

(1961, 1962, 1963, 1966, and 1968) , Szaniawski (1968,

1970, and 1974) , Szaniawski & Wrona (1973) , Corradini & Olivieri (1974) , Jansonius & Craig (1974, 1975) , Boyer

(1975) , Mierzejewski & Mierzejewska (1975) , Mierzejewski

(1978b) , Szaniawski & Gazdzicki (1978) , Edgar (1984) , and

Colbath (1987b) . Most of them have used isolated jaws

together with apparatuses in their taxonomical work. For

the most recent opinions on these questions, see the chap

ter 'Polychaete taxonomy'.

6 Claes F. Bergman

Accounts of fossil annelid jaws from Sweden

Silunan. - Though anne lid jaws are fairly common in the

Silurian strata of Gotland, reports on them are scarce. The

first account of scolecodonts is the well-known one by

Hinde (1882) on annelid remains from the Wenlockian

Hogklint and Slite Beds. Most of his material derived from

the uppermost part, Hogklint unit d, of the Vattenfallet

section (ThorelI & Lindstrom 1885, p. 4; Bergman 1979a).

Lindstrom (1885, pp. 4-5) listed the species described by

Hinde in his list of the Silurian fauna of Gotland, and the

same species also appear in Lindstrom's list of the fossil

faunas of Sweden (1888, pp. 5-6). In a study of the stratig

raphy of the Visby district, Hedstrom (1910) noted the

occurrence of annelid jaws and bristles from the same

section but from lower strata, corresponding to Upper

Visby Beds and Hogklint Beds unit a and b. Bergman

(1979) discussed the reconstruction, abundance and diver

sity ofjawed polychaetes in a study comprising 32 samples

from the uppermost part of the Lower Visby Beds to the

Hogklint Beds unit d of the Vattenfallet section.

In his first work on the geology of Gotland, Hede (1917)

described the fauna from the transitional be ds between the

Wenlockian Slite Mari and Slite Siltstone. His faunal list

induded four of Hinde's annelid taxa. Hede must have

sampled his anne lid jaw collection dose to the second

locality (south of Klintehamn) that Hinde used in his 1882

paper. Hede noted the presence of annelid jaws in a num

ber of localities in his descriptions (1921, 1925, 1927a,

1927b, 1928, 1929, 1933, 1936, 1940).

Angelin, who died in 1876, had prepared a number of

plates to be published in his series Palaeontologica Scandi

naviea, some of which were never published. In one of the

unpublished plates, number 53, a number of more or less

well preserved scolecodonts have been illustrated (figs.

20-26). Later RegneIl (1952, p. 623) made an attempt to

identify the jaws and referred to them as Arabellites cf.

hamatus Hinde, A. cf. contractus Hinde, ?aff. Lumlniconereites

sp., and others. RegnelI used the paper by Hinde (1882) to

make his identifications, and it does not se em unlikely that

Angelin's material derived from one of the same localities

used by Hinde (the locality south of the town wall ofVisby).

This is the uppermost part of the Vattenfallet section

('Pterygotus'marl) on Gotland.

Martinsson (1960) found two assemblages of annelid

jaws from the Hemse Beds, Hemse Mari SE part and Mulde

Mari, respectively, and used the apparatus taxonomy when

he named the material.

Ecological studies of fossil jawed annelids are so far very

rare. However, Laufeld (1975, pp. 804-805) discussed the

abundance of anne!ids and chitinozoans with regard to

benthic marine life zones (sensu Boucot 1975) on Gotland.

Eisenack (1975) described and discussed apparatuses

and isolated elements on material partly from Gotland. He

concentrated on the construction of dentides and possible

resorption and growth lines in mandibles. Later Mierzejew

ski (1984) redescribed one of the apparatuses from the

Silurian of Gotland described by Eisenack (1975).


I have earlier discussed the jawed polychaete fauna from

the Lower Wenlock of Gotland in my first attempts to use

both isolated jaws and apparatuses for a naturai taxonomy

(Bergman 1979a, 1980b, and 1981b). The base for these

reconstructions was a material of more than 13,000 ele

ments from one of the localities. At that time I had not

studied the type specimens involved and therefore I tenta

tively used the same names as used by Hinde. A later paper

(Bergman 1984) de alt with the occurrence of an annelid

jaw that had been transported by a density current.

At present I have studied more than 700 sam pl es from

the Silurian of Gotland (Fig. lA and B, and appendix) ,

representing severai metric tons of different types of lime

stone, marIstone and siltstone. The samples were collected

in order to cover all sedimentary rock types of the island to

indude a total geographical as well as stratigraphical distri

bution. My aim is to study the total jawed annelid fauna of

Gotland, though in this publication only a minor part of

the fauna is presented.

Scolecodonts from Gotland have also been illustrated

and discussed in a popular science book by Brood (1982) ,

who also used the names introduced by Hinde.

Scolecodonts from erratic boulders from Gotland were

pictured by Schallreuter (1982).

Ordovician. - The occurrence of supposed annelid jaws

from Ordovician strata of Sweden has been reported in

various studies, e.g., Wiman (1893) , Westergård (1909) and

Hadding (1913, 1915). The first two accounts only noted

the occurrence of anne!id jaws, but in the latter, Hadding

described and illustrated a num ber of conodont elements

from Scania (Skåne) , be!ieving they were annelid jaws.

These taxa have later been shown to represent conodonts

(Lindstrom 1955). It seems very probable that the observa

tions from the lowermost Ordovician by Wiman and

Westergård also were accounts of conodonts. However,

scolecodonts are present in the Middle and Upper Ordovi

cian on Oland (Yngve Grahn, personal communication).

Erratic boulders. - The source and the age of erratic boulders may be hard to verity. Apart from a few unique specimens

it is probably impossible to state the locality of origin and

exact stratigraphical leve I of the boulders, thus making

them less useful for palaeontological studies (see also chap

ter on taxonomy).

Schallreuter (1982, p. 5) has listed papers on scoleco

donts from erratic boulders of various stratigraphical ages,

some of the boulders treated probably being derived from

the Gotland area.

Geology of Gotland The Silurian strata of Gotland hi de magnificent, three-di

mensionally preserved fossiIs. The sediments, now found

above sea leve!, were forrned in a tropical, shallow and

repeatedly fluctuating intracratonic sea. The palaeoba

thymetry roughly had a NE-SW strike with a mean d/p of

0.15-0.3° (Laufeld 1974a, p. 7) towards the SE (e.g. Mar

tinsson 1958; Agterberg 1958; and Bergman 1984). The


slight dip of the tectonically almost undisturbed layers

exposes a stratigraphical sequence of about 500 m ranging

from Late Llandovery to Late Ludlow in age. The strata are

composed of various types of carbonate-rich sediments

induding biohermal limestone, stratified limestone, oolite,

marlstone and siltstone. The sediments have been depos

ited in subparallel, south-easterly migrating facies beits

with a shore-line to the N to NE.

The sequence has been subdivided into thirteen litho

logical mappable units (Hede 1921 and 1925a; see Figs. 5

to Il for areal extension). These com prise more or less

heterogeneous complexes of strata. Therefore the units

have been referred to as Groups, Formations and Beds by

various authors.

Hede did an impressive piece of field work when he

carefully mapped almost the entire island. The results were

published in eight geological descriptions (1925, 1927a, b,

1928, 1929, 1933, 1936, 1940). Only minor amendments of

Hede's condusions have been made since then (see Lau

feld 1974a and Jeppsson 1983, p. 126). The map descrip

tions are in Swedish, but an English version of his stratigra

phy has been published (Hede 1960, pp. 44-52). Based on

Hede's map descriptions, Laufeld (1974a, pp. 7-13) made

a very useful summary of the stratigraphy and sediments of

Gotland, and he completed Hede's subdivision of the

major units. Awaiting a much needed modern sedimento

logical description of the sediments I refer to Hede's 13

major units as 'Beds' (Laufeld 1974a).

A short summary of the sediments and the Silurian

palaeogeography of Gotland was given by Laufeld & Bassett

(1981). The magnificent bioherms have attracted several

geologists and resulted in special studies (e.g. Manten

1971; Watts 1981). Riding (1981) has given a short review

of the structures of the bioherms.

In his subdivision of the strata, Hede also used the fauna,

with emphasis on some taxa of macrofossils, e.g. Pentamerus

gothlandicus. Later, diachronism between some of the topo

stratigraphical units and the stratigraphical distribution of

fossil assemblages was noted, first by Martinsson (1967).

Other students followed and produced evidence of dia

chronism of severai of the boundaries (e.g., Larsson 1979;

Franzen 1983). Not all suggested diachronisms are ac

cepted by all students (e.g., Jeppsson 1983, pp. 128-129;

Stridsberg 1985, p. 5). It must be pointed out that most of

the stratigraphical work done is based on benthic organ

isms which are more likely to be influenced by the bottom

environments than pelagic organisms are. The conodonts

do not support diachronism of more than a few boundaries

(LennartJeppsson, personal communication, 1986).

Correlation of the Gotland strata with other areas has

been carried out with, for example, graptolites (Hede

1919, 1942) , and ostracodes (Martinsson 1967) , and re

cently Jeppsson (1983) showed that conodonts might be

come the optimal instrument in the correlation of the

Gotland sequence. The relatively limited influence that the

bottom conditions have on the distribution of conodonts,

and their high abundance, makes them very suitable for

correlations.

Silunan pauli nitid polychaetes from Gotland 7

Some students have conduded that in the Gotland area

during the Silurian very shallow water conditions prevailed.

Estimations of the depth have also been made (e.g. Had

ding 1941; Gray et al. 1974). It has become more and more

dear that the maximum water depth was less than earlier

estimated (150-200 m) , probably not more than 90 m

(Riding 1979). During shallower episodes subaerial con di

tions occurred repeatedly (e.g., Watts 1981; Cherns 1982;

Frykman 1985, 1986) .

Stratigraphical and geographical distribution of paulinitids on Gotland

The Silurian paulinitids were most probably carnivorous,

living in burrows in the sediment. The distribution ofjawed

paulinitids is influenced by the conditions on and probably

in the sediment. A large num ber of physical and chemical

parameters are involved. A sedimentological study of the

different samples combined with information of the faunal

distribution would be most interesting. Post-mortem trans

portation of the jaws, bioturbation, and the reworking of

sediment have, of course, changed the original geographi

cal and, probably to a much smaller degree, the strati

graphical distribution.

With the present knowledge of the distribution of the

paulinitids on Gotland and the very fragmentary informa

tion from the surrounding areas, it is impossible to estab

lish the full stratigraphical range of the different taxa.

Thus, the ranges shown he re (Fig. 4) must be considered

as local ranges, controlled by the environment. Thus, even

though some of the lineages seem to show evidence of

diachronism, it is hazardous to use the polychaete distribu

tion as an argument in the discussion of diachronism on

Gotland.

The stratigraphical diagram (Fig. 4) does not account for

the abundance of specimens encountered or the num ber

of localities where the taxon is represented in each unit.

The areal distribution is shown in Figs. 5, 6, 7, 8, 9, 10 and

Il. The abundance of species in some localities of impor

tance is reported in Figs. 2 and 3.

Lower Visby Beds. - These beds are composed of alternating

marlstone and argillaceous limestone. The Lower Visby

Beds were probably deposited at a water depth dose to the

photic zone (Riding 1979) and it belongs to one of the

deepest deposited strata on Gotland (Gray et al. 1974).

Lennart Jeppsson (personal communication, 1987,

manuscript in preparation) has subdivided the Lower Visby

Beds into the subunits b-e for the supramarine exposures

of sediment. Kettnerites (A.) siaelsoeensis is a subspecies

found exdusively in the lowermost unit (b) where it totally

dominates the paulinitid fauna and also the jawed poly

chaete fauna. This subspecies is probably a stenotopic type

with preference for deep water. Kettnerites (K.) abraham

abraham is also found in the Lower Visby Beds unit b, but it

ranges throughout the Visby Beds. K. (K.) martinssonii and

Lanceolatites gracilis are not common in the Lower Visby

Beds, and both have a long range. A characteristic species

of the uppermost unit (e) of Lower Visby Beds is K. (K.) versabilis, induding its varieties. The species is uncommon

8 Claes F. Bergman

.... 'I'I.ANn

""""' :'�." :.

: \ / •.. )

3 1 2

.

) UCk." 1 ( ""."" .. " , • R�nnklint l

,. Nyhamn 4�' 2

51.

• Kambs 5 • Stara Myre 1

• Yxne l

Sjåls6 3 ': Sjlilsl) 1

'( oOacker l IKorPklint 1 G I

Snl1.�k 31 � Snl1ckgl1rdsbaden 1

GUs�:vasvv�� 1

1 } Guslavsvik 2 i �algberget 1 . Tlautet 1

• Blngers Kvarn 2

Sijdra Torg 1 "-Vattenfallsprofllen 1

'Gutevågen 2 Folhngbo 3 ) ........ Vattenfallsprofllen 2

'Foliingbo 1 2 Folhngbo 4 , ' Jakobsberg 1

,Follingbo 2

.] Nygårdsbåckprofllen 1 Slave 1. � Buske 1

• Ygne 2

I Hallbro Slotl 6 • Sion 1

i 1. :.Iuvslajn

• MartJlle 7


Langhammarshammar 1 • "" ...... "" .. " ,f .� f' Au'sv;ken 1 ) Laute'hO,"sv;� 2,

,�aut.'ho,"sv;k 3 rv' Båta 1 SUHhållar 3"-6 iLauter 'I Gamla Hamn 1 ,eGamla Hamn 2

\ ,e Haganas 1 � • Fårt) Skola 1

Strandakersviken ,. Hallagrund l . ... }

Stutsviken 1

H'"udden 1 � V;alms 2', Saxriv 1. \.� ./ - : Nars Stenbrott l

I Nars l "Ar 1

• Svarven l

Halls Huk 1

• Storugns l

• Klinthagen l Stara Banne 2# Stara Banne 3

Gisslause 1.

Slitebrottet l 2 •

Talings 2 •

Hide 1 1·_ • Hide Fiskelage 1 va

OEnholmen 2

2 , Skenalden 1

• TingsHide 1 Cl

• Tomase 1

Gane 2 •

Mo;n., 4 � r. Aistudden

Vike 3 �\ �ke l TjeldershOlm tL·

3 2 8°7-1-

Gothemshammar

Bolvalde l. Fjardinge 1.

Vallslena 2 • • Våsterbjtirs 1

• Suderbys 3 • Kvannvaten l

Bara 1 •

Hijrsne 6" Hijrsne 3

5

Gandarve 1 .. Gandarve 2 Munkebos l •

Vallmyr , .

• Krakfot l • Baju 1

Oslergarnsholm 3 . . .. ? OSlergarnsholm 1 ........ '0 Flale 3 Fjale 2 Garnudden 4 .. �.>.\ I ..... Garnudden 1 Oslergarnsholm 2 '

• NygArds 1 NygArds 2.

.. Godrings l GOdrings 2 • BAlels 1

Garnudden 3. I (;) 4. DjaUPViksuddef. �GrogarnShUvud 1

Traske 1· K�tthammarsvik 1 Herrvik 2 • Båtels 2

' Tule l Hl1lljnge 2 • • Hallinge 1

Slensiu 2. .

.�Vidfl1lle 1"\� �.' .1r.Kuppen 1 2 lambskvle 1 ·�·Angm�ns 2 ) Sut arve 3· i Gulenviks 1. I Gannes 2

Sularve 2 Gannes 3 'Fakle 1

Fig. 1. Sketch maps of the northem and southem part of Gotland, showing the location of the sampled localities and their code names. The size of the dot indicates the number of samples at a locality without regard to sample size or fauna. Base map: the old topographical map sheets 1 : 1 00,000. For descriptions orthe localities, see Appendix (herein).

FOSSILS AND STRATA 25 (1989) Silurian paulinitid polychaetes from Gotland 9

I n:1'"e,

. ) Sysneudd 1 • Tings 1

• Myrsne 1 Gardrungs 1.

MOllbos 2 .. MOllbos 1 Stora Vikare 2.

Ansarve , Gnisvard 1 ,Gnisvård 2

'Oivide 1

o SOjvide 1

• Tjuls 3

Våte 2. • Stenstugårds 1

\ • Pr4stbåtels 2

Våte 1. Stora Tune 1

o Kakhuse 1

Krasse 1· • Vivungs 1

Gyle 1. • Gyle 2

3 re Grynge 1 • MjUklint ( " ... " ..... " ,

• Rudvier 1 • Fågelhammare 3

• Stora Mafrids 2 o Valve 3 • Rågåkre , Lassor' o (

D .Garde 1

• Valbytte 1 • Sigvalde 2

(• Ljugarn �tl Loggarve 2

'Valbytte 2 Valle 1

'LOggarVe 1 GOg8 ,. � t\ '" Valle 2· . Hunninge 1 '\"V) eKlinteberget 1

Robbjåns Kvarn 2 Robbjans Kvarn 3

Tanglings Kvarn 1 • • Tånglings 2 • Botvide 1 Nyan 2 lau Backar 1 •

• �.).. Nabban 2

Svarvare 1 .. ·Mtslner 1 Svarvare 3 .. /. �DIiPPS 2

, Dapps 1 Varsånde 1 • Mulde 2 Klinteenklaven � . • Mulde Tegelbr�k

B�fride 1

• BrotråskkrOken 1

• Autsarve 1 � 0Haugklintar 2 oBofride 2

Valby Bodar 1. • FrlSjel 2

s:;::u��e 1. Amiings 1. • Rangsarve 1

\ .Linde 1

I Gerum��lf;na�:n

2!·t

'\ • Kalldar 1 BIAhl;l.lI 1. o Grymlings 1 Kullands 1 Gardsby ·1

3 "4 . Kalldar 2 Våstlaus 1.

Hallsarve 1 Nyan ,e! • Fie 3 \

Gannor 1 'Gannor 3 • Hallbjanne 1 °Lyrungs 1 Nar 2- �4 C ... J.i eG ' aSSkar

Ondarve 2. 2 � � ,.Kroken 3

H.!lgvlde 1. Nårshamn 2 � (�KrOken 1 2 Glav 1 . Nårshamn 3. • Nars Fyr 1

Burgen 5° -g • Kapelludden 1 Kårne 3.

Askryggen 2 �A Djupvik Gerete 1. • Tjl;l.ngdarve 1 GrlSndalen YQ:- · 2 .'

Trl;l.dgården 1 · Likmide 2 • • Kodings 3

es Burg�n , iJ

Bandlunde 1 o

Malms 1 • Hummelbosholm 1

/'-. \,jrget 1

• • Bjarges 2 Bjarges 1 r

J. Tom'bodarne ,

• Hågur 1 - Sproge 3 . 4

- 5 Sproge

3 2' Snoder

""".�,:::.:. �7 �"'".� '

• Lukse 1

Lilla Hallvards 1 • Snauvalds 1

'Lilla Hallvards 4

Petsarve 1 5. D () Ronnlngs 1. �

.Nisse 1 Kauparve 1 • • Drakarve 1 -Bankvat 1

Grundård 2 ..... Linviken 2

"/ .VaktArd 5 Vak,ård 2 l·taklård 4

Klasård '} oOlsvenne 3

- Uddvide 1

• = Localily wilh ane sample Bodudd , tV

Fig. 1, cont.

Iwo lo four samples five lo eighl samples nine samples or more

-Gisle 1

.Strands 1 Ollajvs 1 •

• Sibbjans 2 • Bottarve 1

• Bottarve 2

Suders 1.

HolmMllar 2.

Barkarveård 1 '�Faludden 2

1.� -'Faludden 1

VOHOlmh."ar , O

.K!ittelviken 5 Husr en 3 ygg !: . V!isterbackar 1

Juves 1 <;>tes 1

Hoburgen 2.3 :"52

_ • Barshageudd 2 ,

.

10 Claes F. Bergman FOSSILS AND STRATA 25 (1989)

LOCALITIES

b

W E N L o C K LOWER VISBY BE DS UPPER VISBY BEDS HOGKLINT BEDS SLITE

b c d e e e a a b b b b c d d d SW a m a r i f mari

� \ � � \ � � � \ \ \ , � � � � � , � � � � � � \ � er t-iS' & $ (\)

o:: "� "'..-: IQ IQ .... oS' "'...:: "..-:: "'..-:: "'..-:: 1.. 19., "...-:: "..-:: "' .... /..\9 l.Y t-" -4- "'0 ? �.,: ? l.Y .., l.Y ? 1I'6� �� 'l: iS'.f..-: II'+/. iS'I+/. �� /,-: �/,-: q>-:.: ? ? �'l: <$)'l: �'l: ? :':\9 � � � � � � � � � % � � � � � � � ., � � % % % v ? /' � % % % % % % % ? � �

�� "? &.., / / ,)' / ? ? ? ? ? �� ""� /' , . " " " ' " e

,. ?� $;<, 7 9 " 1 2 75102 81 " 6 8 2 " 8 7 9- 0 0 7 9"42 7 6 " 6 7 6" 8 8 1 -42 7 6 " 9 76"1 0 7 3 -57 7 0 " 2 0 70" 6 7 0 " 5 70" 2 84"71 84" 3 RM RM RH 9"46 8H6 84"75 84-67 83-36

':S�" L J C B L J L J L J L J L J L J L J C S C B L J L J L J L J L J L J L J LJ L J LJ L J L J I>

SPECIES 4 403 5 0 0 4270 2 820 25 6 8 2 592 2000 2000 3 650 506 500 5 4 3 3 670 2 5 00 500 2500 2 25 0 1 800 '3000 930 5 ' 6 2 5 9 0

r M I 64 7

K. (A.) siaelsoeensis I Ml 6 9 2

r M I l 5 4 3

I M l ! 3 8 1

r M I 6 2 2 , 7 9 8 , 1 7 3 2 X 2

K. ( K . ) a b r a h a m I M I 7 1 , 7 9 9 3 1 6 3 3 X a b r a h a m r H I l 4 1 0 3 8 12 1 0 4 , 9 3 3 X "' I M l ! 1 2 7 2 ' 6 9 6 4 ' 8 4 , X

r M I 1 8 9 8 2 3 2 6 1 1 1 3 7 5 3 9 3 3 3 2 5

K. ( K . ) m a r t i n s s o n i i I M I 1 7 1 1 7 2 2 4 1 , O 9 7 3 2 4 4 2 2 3 1

,. H l I 3 , 1 9 1 2 ' 8 5 3 3 7 2 7 8 , 60 , 5 1 1 , 1 8

I M I l 1 3 9 ,. 1 ' 8 6 1 3 6 4 37 9 6 4 3 1 , 5 r M I 1 , 3 ' 4 5 7 , 4 4 1 4

L. g r a c i l i s I M I , 1 2 8 1 2 8 5 3

r M I I 1 2 1 1 3 3 7 1 2 6 2 3

I M l ! , 5 4 4 , 1 1 4 1 2

r M I ' 8 9 2 5

K . ( K J v e r s a b i l i s I M Il 2 2 6 1 6

r M I l 1 8 6 5

I M l ! ' 8 5 3 3

r M I 4 2 1 0 1 0 6 8 ' 9 3

K . ( K . ) a b r a h a m i s a a e I M l 1 8 1 2 2 ' 0 1 0 9 9 4 1 2

r M I t 1 2 2 2 3 6 9 3 8 4 1 ' 4 3 7

I M I l 9 2 , 9 1 3 8 2 1 0 4 ' 5 3 3

r M I 1 1 1 6 2 0 5 9 5 3 7 , 3 2 9 5

K. ( A J m i c r o d e n t a t u s I M I , 1 5 1 2 2 4 5 3 , 3 3 2 8 2

r M l I 3 , O 1 2 2 4 5 , , 2 2 5 5 4 I M I l , 2 6 9 1 3 , 2 2 2

r H l 1 3 2 4 8 4 2 2

H. g l a d i a t u s I H I ' 6 1 2 6 ,

r H U 1 0 6 3 , 1

1 M I l 7 2 0 1

r H l 1 8 3

H. a n g u s t u s I M I 1 1 6 3 9 1

r M I l , 3 2 6 1 , I H I l 4 3 2 1

r M l 6

K. ( A . ) s i s y p h i var. valle I M l 6

r M I l 2

I M l l 2

r M l

K . ( K . ) b a n k v a e t e n s i s I H I

.. H I l 3

I M I l

r M l 3 1

K . ( K .) j a c o b i I M I 3 , .. H I l 3 7

I H l ! 2 3

r M l 2

K . ( A J s i s y p h i s i s y p h i I M I 5

r H l I 1

I M l ! ,

T O T A l 7 1

7 '

5 7

3 9 6 0

5 9

7 6

7 9

' 8 4

1 3 6

2 3 0

1 5 O 4 5

3 ,

4 '

2 4

3 4

3 5

2 9

2 9

6 2

1 7

1 2 8

1 1 5

87

61

6 5

2 9 7 1

4 4

2 1

2 8

4 9

5 6

3 4

3 7

6

6

2

2

3

4

4

1 0

5

2

5 1

1

2 4 9 5 �

Fig. 2. Distribution of paulinitid jaw elements in a few �lected samples from the lower part of the Gotland strata i.e. Lower and Upper Visby Beds, Hogklint Beds (no identifiable elements encountered from the Tofta Beds), and part of the Slite Beds (the uppermost part of the Slite Beds is reported in Fig. 3). Only the four main elements have been included in the count. The letter X denotes that the identification of elements is doubtful or that it has been impossible to separate the homologous elements of the different taxa.

outside this unit, but it is found in unit d and b and from

below unit b (a LennartJeppsson sample, collected about

14 m below water level at Ireviken, not included in the

locality list).

Upper Visby Beds. - Lithology similar to that of the Lower

Visby Beds, but with the first sign of reef formation (e.g.

tabulate-dominated mounds). The U pper Visby Beds were

probably deposited in shallower water than the Lower Visby

Beds.

The fauna is consistent within these strata. Kettnerites (K.) abraham abraham, K. (K.) martinssonii, and Lanceolatites grac

ilis and L. gracilis var. visby occur concurrently through the

Upper Visby Beds. Their ranges are not exclusive for these

strata, but their co-occurrence is typical of the U pper Visby

Beds. Lanceolatites gracilis and L. gracilis var. visby are usually

found in mari, which has been deposited in water of mod

erate to greater depth on Gotland. One subspecies which

can be found throughout the succeeding parts of the Got

land sequence appears, viz. Kettnerites (A. ) microdentatus.

Kettnerites (K.) versabilis disappears in the lower part of the

Upper Visby Beds.

Hiigklint Beds. - The most conspicuous rock types are large

bioherms and argillaceous limes tones intercalated with

marIstone. The water energy varied considerably.

FOSSILS AND STRATA 25 (1989) Silunan paulinitid polychaetes from Gotland Il

WENLOCK LUDLOW

SLITE HALLA MULDE KLINTEBERG HEMSE EKE BU. HAMRA mari 9 Pg Pg b c I u a a e b NW NW SE SE SE b b b

LOCALITIES : ~ o~ :':>$ ~ l-t9 ~ <SI.,: ~ ~ ~ d'~ dI~ "o.JI (SI -2 "'o.JI O' O' ~ ~ ~"? oS' ~I' 0$1,...

~ ~ ~..... "'of oSI...~ .... ~ % ~ (5\ ~ ~ ~ ? .

~ .,~

~ '. ' e '" ,~~ '"." e' ,", 'e ' "

.;<)~ l-l~ I73-2 84-288l-4 66141 71-8181-7577-475-2282-7 82-l773-l0 75-3 77-7 83-5071-4082-1482-31 l-ISO 1-1518215 H99 83-12 76-19 82-l082-34 ' <; LJ ' LJ LJ C8 SL LJ C8 LJ C8 ca C8 LJ CB CB LJ LJ ca LJ LJ LJ CB LJ LJ ca CB ca TOTAL

<> .

SPECIES 500 500 290019000 45 500 500 590 550 550 2249 2520 3500 9770 5250 500 1829 573 5300 677

rMI 3 l 7 14 5 7 l 25 12 12 5 4 12 l l 10 6 3 12 2 145

K . (KJ martins50nii IMI l 8 31 5 11 l 41 17 15 7 8 7 2 2 8 7 2 10 18 1 rMU l 3 II 14 11 2 17 18 15 14 4 9 5 l 8 l l 27 195 IMIl 2 2 2 25 12 8 3 16 8 9 10 l 12 2 7 9 l 8 15 l 15 3 rMI 4 67 9 4 8 2 9 6 2 2 7 10 13 0

K . (AJ mi c rodentatu s IMI 2 63 10 4 6 2 8 10 l 7 19 13 2 rMIl l 41 X X X 11 X 8 3 64 I MIl l lO X 4 3 X 5 X 7 2 5 2 rMI 11 5 26 8 20 157 l 228

K . (K.) bankvaetensis IMI 10 8 l5 14 12 172 l 252 rMIl 18 X 8 17 9 20 164 6 242 IMU 16 X 9 Il 4 10 133 2 187

rMI 33 16 l 7 6 9 72 K. (A .) s isy phi sisy phi IMI l lO 7 6 4 8 56

rMI! X 13 X 22 X 6 4 12 67 IMII 6 X 9 X 3 5 6 29 rMI 9 13 20 6 l 27 76

K. (K.) huberti I MI 9 14 23 l l l 7 2 29 87 rMIl 6 14 7 l 3 2 33 66 I MIl 3 5 15 3 21 47 rMI 3 7 10

K. (A .) fjaelens is I MI 2 10 12 rM Il 2 8 10 I MU O 7 7 rMI 3 l 18 l 23

G. slitensis IMI l 18 19 rMIl 6 l 31 l 45 I MIl 2 l 19 22

rMI l l 2 4

H. gladia tu s I MI l l rMIl IMIl rMI 41 41

K . (AJ sis y phi var. valle IMI 42 42 rMIl X 50 50 IMIl X 32 2 34 rMI l 10 22 87 l 121

K . (K.) polonensis I MI 9 11 17 2 77 6 122 r MIl 2 7 12 l 61 2 3 88 I MIl 2 4 13 l l 54 4 l 80

rMI l l

H. angustus IMI rMIl l l IMIl rMI 6 10 36 l 53

L. gracilis I MI 7 l 49 l l 59 rM Il 6 10 l 45 l 63 I MIl 4 9 34 2 49

rMI 40 4 4 48

K . (K .) burgensis I MI 52 52 rMIl 65 2 2 69 I MIl 19 l l 41

r MI 8 23 13 44 K. (A.) s iSyphi I MI 10 18 15 43

klasaardensis rHII 3 19 20 42 IMIl 5 14 15 34

rMI . 13 13

H. naeren s i s IMI l 16 19

rHIl l 4 5 I MIl 2 4 6

rMI l l

Lanceolatites sp . A IMI 4 4 rHlI 5 5

IMIl 2 2

3848 '---

Fig. 3. Distribution of paulinitid jaw elements in a few selected samples from the upper Slite Beds to the Hamra Beds. Only the four main elements have be en included in the count. The letter X deno tes that the identification of the elements is doubtful or that it has been impossible to separate the homologous elemen ts of the different taxa.

The evolution from K. (K.) . abraham abraham into the

younger subspecies K. (K.) abraham isaacoccurs within the Hogklint Beds, unit b (possibly also a), both subspecies being characteristic and easily identified taxa_ K. (K.) abra

ham isaac continues through the Hogklint Beds and is not firmly identified outside these strata. On ly one occurrence

is recorded from a sample of younger strata in the Halla Beds. The identification, based on a few poorly preserved jaws, is doubtful [K. (K.) aff. abraham isaacl . The genus Hindenites appears in the Hogklint Beds with two lineages, H gladiatus and Hangustus). The two speeies are of ten but not always found together. The Hogklint Beds are the only

12 Claes F. Bergman

strata on Gotland where H. gladiatus and H. angustus are

fairly common. They probably Iived in shallower water in

the Gotland area. Kettnerites ( K. ) martinssonii is frequently

found in every sample and K. (A.) microdentatus is also fairly

common. Two new lineages appear, K. ( K. ) bankvaetensis

and K. ( K. ) polonensis, but only a few jaws are recorded, and

their identification is uncertain, based mainly on the sec

ond maxillaryjaws (MII's) .

Tofta Beds. - Only an unidentifiable jaw fragment has been

encountered in the Tofta Beds. Unfavourable living condi

tions, high water turbulence, coupled with bad preserva

tion conditions are probable explanations of the lack of

fauna. The same environment seems to have persisted

during the formation of the lowermost part of the Slite

Beds.

Slite Beds. - The Slite Beds are composed of two main

Iitho-faeies: limestones (typical of the earliest units) and

marIstones (typical of the succeeding units) . There are a

number of exceptions, however. The strata indude a rich

variety of sediments such as siltstone, reefal limestone,

stratified limestone and marIstone. The earliest units, a to

e, do not show any distinct faunal characters beside a very

low abundance coupled with low diversity. A maximum of

only two species are found in each unit. Kettnerites ( K. )

martinssonii is a typical representative of the fauna but must

be regarded as an eurytopic speeies on Gotland. The spe

eies can be found both in typical mari levels and in lime

stone Ievels more or less all over the island.

A dramatic change of the fauna, probably inflicted by an

environmental change, occurs in the undifferentiated part

of the Slite MarI. A rich and varied fauna is encountered,

particularly in the Slite MarI unit with Pentamerus gothland

icus. Typical speeies are Kettnerites (K.) jacobi and Gotlandites

slitensis. K. ( K. ) jacobi is a conspicuous speeies, of ten with

large jaws. It has a short stratigraphical range (Slite Mari,

undifferentiated part to Lerberget Mari) on Gotland and is

typical of unit g. Gotlandites slitensis is typical of the Slite

Mari unit Pentamerus gothlandicus but ranges from the Slite

Mari, undifferentiated part (Hide l) , to Halla Beds, undif

ferentiated part (Gandarve l , where only two jaws are

encountered) . Kettnerites ( K. ) huberti, K. ( K. ) bankvaetensis,

and K. (K. ) polonensis lineages, all with long stratigraphical

ranges, are first recorded with certainty in the Slite Beds.

The latter two have been found in earlier strata but there

on ly tentatively identified.

Halla Beds. - The strata consist of argillaceous limestone,

small mounds and oolite, all deposited in a shallow-water

environment.

The fauna in the Halla Beds has normally a fairly low

diversity and abundance with only a few species per sample.

K. ( K. ) bankvaetensis and K. ( K. ) polonensis become more

common than in the Slite Beds. Together with K. ( K. )

martinssonii they totally dominate the fauna. K. (A.) fjael

ensis appears.

Mulde Beds. - These are argillaceous limestone alternating

with marIstone. The strata are more or less equivalent to


the Halla Beds in time. The water energy was low but the

water depth was probably fairly shallow.

The fauna is similar to that of the Halla Beds, but Kettner

ites ( K. ) martinssonii becomes most conspicuous, with the

largejaws of a new variety [ K. (K. ) martinssonii var. muldel .

This speeies dominates the fauna, together with K. (A.)

sisyphi. The morphology of the latter speeies is variable,

between the localities and of ten also within a sample. The

latest recorded occurrence of Hindenites gladiatus and K.

( K. ) polonensis var. gandarve is recorded from these Beds.

Klinteberg Beds. - The lithology varies from well-washed

white limestones to argillaceous limestones. Large bio

herms are also typical.

The poor recovery of paulinitids from these be ds is prob

ably due to unfavourable Iiving conditions and poor pres

ervation. With few exceptions, the samples have yielded

only a few fragmented jaws, but occasional extremely well

preserved jaws are encountered. In the NE part (e.g. Got

hemshammar) fairly rich faun as are recorded. A typical

Klinteberg fauna of the eastern type is dominated by Ket

tnerites bankvaetensis with K. ( K. ) polonensis and K. ( K. )

martinssonii as normal accessory elements of the fauna. K.

( K. ) bankvaetensis is rare in the western area of the

Klinteberg Beds. It is also rare in both the other younger

and the older strata on the western side of Gotland (viz. in

the more argillaceous sediments) . The latest occurrence of

the Hindenites angustus lineage is recorded from the lower

middle part of the Klinteberg Beds.

Hemse Beds. - The Iithology is similar to that of the Slite

Beds, with a large number of different types.

The rich and variable fauna is also reminiscent of the

younger Slite fauna, particularly in some of the marly areas

of the latter. Kettnerites ( K. ) huberti, earlier recorded from

only two localities in the Slite Beds, is very common and a

typical speeies in the Hemse Beds, except in the Hemse

MarI, NW part. Kettnerites ( K. ) polonensis is another common

speeies, particularly in sediments with a lower proportion

of argillaceous partides. K. (A.) sisyphi klasaardensis and K.

( K. ) burgensis ·are alm ost endemic of the highly argillaceous

sediments in the Hemse Mari SE and Mari Top. Both these

lineages have a short range on Gotland. They appear in the

Hemse MarI, NW part but very sparsely. Typically K. (A.)

sisyphi klasaardensis dominates the paulinitid fauna in the

Hemse Mari, SE part, particularly in the southeasternmost

part along the present coast-line. K. ( K. ) burgensis seems to

disappear in the latest Hemse Mari dose to the boundary

with the Eke Beds, while K. (A.) sisyphi klasaardensis contin

ues through the boundary strata. K. (A. ) microdentatus is

Fig. 4. Stratigraphical distribution of pauliI!itid jawed polychaetes on Gotland. Solid squares (.) represent fully identified taxa, solid cireles (e) less confiden tly assigned ones. Open squares (O ) represent identified taxa from localities with an imprecise stratigraphical position (e.g. lower-middle). The stratigraphical column does not reflect the true stratigraphical succession;Jeppsson (1983, Fig. 2) gives a more correct stratigraphical column but the coexisting units make it inappropriate to use in this connexion. The stratigraphical column is modified from Laufeld 1974a, Fig. 77, and

Jeppsson 1983, Fig. 2.

LLAN

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14 Claes F. Bergman FOSSILS AND STRATA 25 (1989)

G O T L A N D H o g k l i nt B e d s

T o f t a B e d s

{) Q M u l d e

B e d s

B u r g s v i k

Low e r & U p p e r

V i s b y B e d s

. *

.

* '/

,*� •.

(! D

> ,ro "* A

-.H a l l a B e d s

�./

Y· ' .

K l i n t e b e r g B e d s

j,' ; '" �. . � .. �. . ;;' �.J'\ å o

.

4 ...

r;') ��m s e B e d s

('

0<)

B e d s

A '. , � . . N . ' . •

B e d s

1 0 k m .... ====;;;;;1'

I

* K e t t n e r i t e s ( K . ) a b r a h a m a b r a h a m

• K . ( K . ) a b r a h a m i s a a c

* K . ( K . ) j a c o b i

• K . ( K . ) b u r g e n s i s

Fig. 5 . Sketch map with the investigated localities represented by small dots and other symbols (see legend) indicating the identified taxa. Stratigraphical ranges: Kettnentes (K) abraham abraham, Lower Visby Beds, unit b, to Hogklint Beds, unit b. K (K) abraham isaac, Hogklint Beds, unit b, to Hogklint Beds, southwestern facies. K (K) jacobi, Slite Beds, Slite Mari, undifferentiated part, to Lerberget MarI. K (K) lrurgensis, Hemse Beds, from the Hemse Mari, NW part, to the Hemse Mari, uppermost part.

FOSSILS AND STRATA 25 (1989) Silurian paulinitid polychaetes from Gotland 15

G O T L A N D r· o �.(;

C) M u l d e B e d s

o

/ ø<:::7 }) H o g k l i n t B e d s .) " �. /\ r'"��

oJ.J � >Jr , T o f t a B e d s fr " . *' V

il \} S l i t e B e d s o� o �Ha"a B e d , �� /

Oo

.. l, ) V o. o •


h· ' . � �. . .. � 7�" · �.f'I\ ... /. " � o .>"j , ,"'-. o � �y ( H e m s e B e d s

)� � . ,. , • M .: . , . <:;, . .

; ,,- ' A N I

• K etlner i tes ( K.) v e r s a b i l i s

* Got lan d i t e s s l i t e n s i s

• K e t t n e r i t e s ( K . ) h u b e r t i

Burgsvik B e d s

B e d s

1 0 km ==� ..... ��..JI

Fig. 6. Sketch map with the investigated localities represented by small dots and other symbols (see legend) indicating the identified taxa. Stratigraphical ranges: Kettnerites (K) versamiis, Lower Visby Beds, unit b, to the lowest part of the Upper Visby Beds. Gotlandites stitensis, Slite Beds, Slite Mari, undifferentiated part, to Halla Beds. K (K) huberti, Slite Beds, Slite Mari, undifferentiated part, to Hamra Beds, unit c.

16 Claes F. Bergman

G O T L A N D

T o f t a B e d s

L ow e r & U p p e r

V i s b y B e d s

• � M u l d e

B e d s

B u r g s v i k B e d s B e d s

o 1 0 k m Ib=�==dl



A N I

• K e t t n e r i t e s ( A . ) siaelsoeensis

... K . (K.l b a n k v a e t e n s i s

* K . ( A . ) s i s y p h i s i s y p h i

* K . (A . ) s isyph i v a r . val le

• K . (A.) s isyph i k l a s a ardens is

Fig. 7. Sketch map with the investigated localities represented by small dots and other symbols (see legend) indicating the identified taxa. Stratigraphical ranges: Kettnerites (A. ) siaelsoeensis, Lower Visby Beds, unit b. K (K) bankvaetensis, fully identified from Slite Beds, unit d, to Sundre Beds. K (A. ) sisyphi sisyphi, Lower Visby Beds, unit b, to Hemse Beds, Hemse Mari, SE part. K (A. ) sisyphi var. valle, fully identified from the Slite Beds, unit Pentamerus gothlandicus, and probably ranging to the Halla Beds, unit b. K (A. ) sisyphi klasaardensis, Hemse Beds, Hemse Mari, SE part, to Eke Beds, lowest part.

FOSSILS AND STRA TA 25 (1989)

G O T L A N D

T o f t a B e d s

O

� Q M u l d e

B e d s

L o w e r & U p p e r

V i s b y B e d s

• •

...

...

..

• •

� . . /."

· 1 · •

...

c1 c:>

o Ll

o

Silunan paulinitid polychaetes from Gotland 17

H o g k l i n t .- . f\-/ B e d s • � �

� o

S l i t e B e d s


";j:. : .�) CV

Fn �e m s e B e d s

od A Q,

N I

• Kettner i tes ( A . l m i c r o d e n t a t u s

... K . ( A . l f j a e l e n s i s

* L a n g e i t e s g l a b e r

Fig. 8. Sketch map with the investigated localities represented by small dots and other symbols (see legend) indicating the identified taxa. Stratigraphical ranges: Kettnentes (A. ) microdentatus, Lower Visby Beds, unit b, to Hamra Beds, unit b. K (A. ) fjaelensis, fully identified from the Halla Beds to the Lower Eke Beds. Langeites glaber, Hemse Beds, Hemse Mari, SE part, to Sundre Beds.

18 Claes F. Bergman FOSSILS AND STRATA 25 ( 1989)

G O T L A N D

T o f t a B e d s

L o w e r & U p p e r

V i s b y B e d s

B u r g s v i k B e d s

.. . ' \::; � V S l i t e B e d s

•

D

> :f' H a l l a B e d s

,. ff/I . . dv •

• K l i nt e b e r g B e d s

� . . . �.� . ,-.

n' ,,; r . ��m s e B e d s

(' A N I

• K e t t n e r i t e s ( K . l m a r t i n s s o n i i

o 1 0 k m \;;;;;;;;;;====='

Fig. 9. Sketch map with the investigated localities represented by small dots and the filled square representing Kettnmtes (K.l martinssonii, ranging from the Lower Visby Beds, unit b , to the Hamra Beds, unit b.


d

G OT L A N D

T o f t a B e d s

L o w e r & UPPlfeit •

V i s b y B e d s I/! , ! �. it #. " t ' . . o

tJ M u l d e B e d s

-.

"'. Yr

,.) , .

. . .�

-: . •• .

.

B u r g s v i k

Silunan paulinitid polychaetes from Gotland 1 9

(.

l'·

/J . J ·d o

o o

B e d s

1 0 k m ,=====;;;:JI

B e d s

B e d s

B e d s


A N I

• K ettn e r i t e s ( K . ) p o l o n e n s i s

Fig. 10. Sketch map with the investigated localities represented by small dots and the filled triangle representing Kettnerites (K.) polonensis, typical populations of which range from the Slite Beds, Slite Mari, undifferentiated part, to the Sundre Beds.

20 Claes F. Bergman FOSSILS AND STRA TA 25 ( 1 989)

G O T L A N D

o

T o f t a B e d s

{:� S l i t e B e d s

© Q M u l d e

B e d s

B u r g s v i k

> A( �o H a l l a B e d s

... /--. . - . ;;jO .

4! 'O o o.


�o - 2000 �o o 00 ;;�

o �of'\ Ll .lo � o o �) • o il

io o 'ft ° f"� m s e B e d s

'3 ° . , , ' ./ 0 0 o

•

. o� o : •

B e d s

B e d s

B e d s

1 0 k m I

•

...

T

•

O

H i nd e n i t e s angustus

H . g l a d i at u s

H . n a e r e n s i s

L a n c e o l at i tes g r a c i l i s

L a n c e o l a t i t e s s p . A

A N I

Fig. 11. Sketch map with the investigated localities represented by small dots and other symbols (see legend) indicating the identified taxa. Stratigraphical ranges: Hindenites angustus, Hiigklint Beds, unit c, to Klinteberg Beds, lower part. H. gladiatus, Hiigklint B·eds, unit b, to Mulde Beds, upper part. H. naerensis, Burgsvik Beds, lower part. Lanceolatites gracilis, Lower Visby Beds, unit b, to Eke Beds, middle part. L. sp. A, Hamra Beds, unit b.

FOSSILS AND STRATA 25 ( 1 989)

another example of a species which is found in sediments

with a high argillaceous con tent. A very rare form, Langeites

glaber, is recorded from two localities in the Hemse Mari,

SE part, with only a few specimens. Lanceolatites gracilis is

found in both the NW and SE parts of the Hemse MarI . In

the localities where it is encountered it is of ten fairly abun

dant.

Eke Beds. - These are argillaceous limestones and mud

stones, in places rich in calcareous algae, and in the north

eastem part rich in small bioherms.

Paulinitids are normally less common in the Eke Beds

than in the H�mse Beds. The change in the composition of

the paulinitid fauna through the Hemse-Eke boundary is

gradual. Kettnerites ( K. ) burgensis does not reach the base of

the Eke Beds while K. (A. ) sisyphi klasaardensis and K. (A. ) fjaelensis pass through the boundary beds but disappear in

Fig. 12 (p. 22) . Hypothetic phylogenetic relationships between Gotland paulinitids. The connecting lines show probable or, if broken, possible relationships. Note that due to lack of space it has not been possible to place all the apparatuses in relation to each other in their proper stratigraphical orientation. The size of the drawings reflects roughly the mean size of thejaws (magnification

about x20) . However, the smallestjaws have been enlarged and the largest jaws have been reduced in size in comparison with the medium-sized jaws. A comparison of size is also possible between Figs. 12 and 13 . The jaws of Fig. 14 are, however, reproduced at a larger magnification (about x30) than those in Figs. 1 2 and 1 3, due to the minute size of the jaws of some of the taxa included in that figure . Screening patterns represent the lithologies (bricks=limestone; broken ruling=marlstone) in which the illustrated speci

mens are found. DA. Kettnentes (K ) martinssonii, left and right MI and MIl, Vattenfallsprofilen l, Hogklint Beds, unit b, sample 70-6LJ. DB. K (K) martinssonii, left and right MI and MIl, Valle 2 , Slite Beds, Pentamerus gothlandicus Beds, sample 66-l 45SL. DC. K (K) martinssonii var. mulde, left and right M I and MIl jaws from adult specimens, Snoder 2, Hemse Beds, Hemse Mari, NW part, sample 82-l 4CB. OD. K (K) martinssonii, left and right MI and MIl,

Snoder 2, Hemse Beds, Hemse Mari, NW part, sample 82-l 4CB. D E. K (K) martinssonii, left and right MI and MIl, Sigvalde 2 ,

Hemse Beds, lower-middle part, sample 7l-1 1 5LJ. O F. K (K) martinssoni� left and right MI and left MIl, Glasskar 3, Burgsvik Beds, lowest bed, sample 82-l 8CB, right MIl, Glasskar l , Burgsvik

Beds, lowest part, sample 82-l5CB. O G. K (K) martinssonii, left MI and right MIl, Kauparve l , Hamra Beds, lower-middle part, sample 76-l 3CB, right MI and left MIl, Narshamn 2, Burgsvik Beds, lower part(? ) , 83-l 2LJ. OH. K (A. ) siaelsoeensis, left and right MI and MIl, Sjalso l, Lower Visby Beds unit b, sample 79-l 2LJ. DI . K (A.) microdentatus, left and right MI and MIl, Håftingsklint l , Upper Visby Beds, sample 76-9CB. DJ. K (A. ) cf. microdentatus, left and right MI and MIl, Vattenfallsprofilen l , Hogklint Beds, unit b, sample 70-6LJ. D K K (A. ) sisyphi var. valle, left and right MI and MIl, Valle 2 , Slite Beds, Pentamerus gothlandicus Beds, sample 66-l 45SL. D L. K (A.) sisyphi sisyphi, left and right MI and MIl, Dapps

2, Mulde Beds, upper part, sample 82-37CB. DM. K (A.) sisyphi sisyphi, left and right MI and MIl, Varsande, Mulde Beds, lowest part, sample 75-22CB. ON. K (A.) sisyphi klasaardensis, left and right MI and MIl, Vaktård 4, Hemse Beds, Hemse Mari, SE part, sample 81-35LJ. DO. K (A.) sisyphi sisyphi, left and right MI and MIl, Snoder 2, Hemse Beds, Hemse Mari, NW part, sample 82-1 4CB. DP. K (A.) fjaelensis, left and right MI and MIl, Fjale 3, Klinteberg Beds, unit e , sample 77-7CB. D Q. K (A.) microdentatus, left and right MI and MIl, Stave l , Slite Beds, Slite Mari, 75-1 1 CB. D R. K (A.) microdentatus, left and right MI, Narshamn 2, Burgsvik Beds, lower part ( ? ) , sample 82-12LJ.


the basal part of the Eke Beds. K. ( K. ) bankvaetensis, K. huberti, and K. (K. ) polonensis are the dominating species in

the Eke Beds.

Burgsvik Beds. - In the south, the Burgsvik Beds are dominated by clastic sediment (sand-, silt-, and claystone) and oolites to pisoolites, all with a low abundance ofjaws. In the northeastem part the extension of the Burgsvik Beds is composed of an arenaceous limestone and biohermal limestone.

A polychaete fauna with a high abundance but fairly low

diversity is encountered in the northeastem part of the

areal extension of this unit. The most dominating species

in the northeastem part of the boundary between the Eke

Beds and the Burgsvik Beds is Kettnerites ( K. ) polonensis. The

Burgsvik Beds, lower part, is characterized by the conspic

uous Hindenites naerensis. This species is, however, fairly

uncommon. Kettnerites (A. ) microdentatus, which is usually

found only in low numbers on Gotland, is here among the

more common species, together with the eurytopic K. (K. ) martinssonii.

Hamra Beds. - The argillaceous limestone intercalated with

marIstone characterizes the middle unit b. Biohermal lime

stone is found in the upper part of the Hamra Beds.

Fig. 13 (p. 23) . Hypothetic phylogenetic relationships between Gotland paulinitids. See Fig. 1 2 for explanation. DA. Kettnentes (K ) polonensis var. gandarve, left and right M I and MIl, Gothem� hammar 7, Halla Beds, unit c, sample 77-45LJ. DB. K (K) polonensis, left and right MI and MIl, Glasskar 3, Eke Beds, sample 82-1 8CB. DC. K (K) polonensis, left and right MI, Glasskar l , Eke

Beds, sample 82-15CB. OD . K (K) polonensis, left and right MI and MIl, Faludden 2, Hamra Beds, unit c, sample 76-1 6CB. DE. K (K) polonensis, left and right M I and MIl, Sibbjans 2 , Hamra Beds, unit b, sample 82-32LJ. O F. K (K) polonensis var. sjaustre, left and right MI and MIl, Sjaustrehammar l, Hemse Beds, unit d, sample 82-1 9LJ. O G. K (K) sp. A, left and right MI and MIl, Likmide 2, Hemse Beds, Hemse Mari SE part, sample 82-28LJ. OH. K (K) burgensis, left and right MI and MIl, Vastlaus l , Hemse Beds, Hemse Mari, SE part, sample 82-31LJ. DI . K (K) bankvaetensis, left and right MI and MIl, Vike 2, Slite Beds, unit Pentamerus gothlandicus, sample 834CB. DJ. K (K) bankvaetensis, left and right MI and MlI, Mollbos l, Halla Beds, unit b, sample 77-28LJ. DK K (K) bankvaetensis, left and right MI and MIl, Fjardinge l , Klinteberg Beds, unit b, sample 77-SCB. D L. K (K) bankvaetensis, left and right MI and MIl, Bankvat l, Hamra Beds, unit b, sample 82-30CB. D M. K (K) bankvaetensis, left and right MI and MIl, Kauparve l , Hamra Beds, lower-middle part, sample 76-1 6CB. ON. K (K) abraham abraham, finely denticulated, left MI, Nygårdsbackprofilen l , Lower Visby Beds, unit e, sample 79-42LJ, right MI, Lickershamn 2, Lower Visby Beds, unit e, or lowest part of Upper Visby Beds, sample 73-53LJ. DO. K (K) abraham abraham, left and right MI and MIl, Buskt; : l , Lower Visby Beds, unit e , sample 79-40LJ. DP. K (K) abraham isaac, left and right MI and MIl, Vattenfallsprofilen l , Hogklint Beds, unit b, sample 70-6LJ. D Q. K (K) jacobi, left and right MI and MIl, Slitebrottet 2, Slite Beds, unit g, sample 83-31 LJ. D R. K (K) huberti, left and right MI and MIl, Hide l , Slite Beds, Slite Mari, sample 73-2LJ. DS. K (K) huberti, slender type, left and right MI and MIl, Vaktård 4, Hemse Beds, Hemse Mari, SE part, sample 81-35LJ. DT. K (K) huberti, straight type, left and right MI and MIl, Bankvat l, Hamra Beds, unit b, sample 82-34CB. DU. K (K) huberti, coarse denticulated type, left and right M I , Bankvat l , Hamra Beds, unit b, sample 8l-39LJ, left and right MIl, Bankvat l , Hamra Beds, unit b , sample 82-32CB.

22 Claes F. Bergman FOSSILS AND STRATA 25 ( 1 989)

K . ( K .) martinssoni i l i n e a g e

K . ( A . ) m i c r o d e ntatus l i n e a g e

o

...J

o

...J

u

o

...J

z

w

Fig. 12 (caption on p. 21).

D STRATA 25 ( 1 989) FOSSILS AN

K. . (K . l po lonens is

. on p. 2 1 ) . . 13 (captlOn Flg.

l ineage

f..om Gotland 23 l · 'tid polychaetes J' Silurian pau mz

K. (K . ) bankvaetens is l i neage

K . (K . ) hu bert i l i neage

24 Claes F. Bergman

o

L a n c e o l a t i t e s sp . A

l i n e a g e

I Lanceo lat i tes g r a c i l i s l i n e a g e

H . n a e r e n s i s l i n e a g e


Lange i tes g laber l i ne a g e

K . ( K . ) v e r s a b i l i s l i n e a g e


Only unit b yields polychaete jaws in any gre at amount.

The jaws are of ten very large, and from some samples (e .g.

Bankvat l) the fauna is totally dominated by one species.

Severai lineages seem to disappear in this unit. This is

probably only an artifact, because the Hamra Beds, unit c,

and the Sund re Beds are fairly poor in annelid jaws. Thus,

the lineages may continue into younger strata exposed

south east of Gotland. One new species, Lanceolatites sp. A,

appears and disappears in the Hamra Beds, unit b.

Sundre Beds. - Mostly crinoidal and biohermal limestones.

Very few jaws, of ten fragmented, are encountered in these

strata, and those found are of ten badly preserved. Shallow,

high ly turbulent water with unfavourable bottom condi

tions seems to be the explanation that the fauna is so

limited. Very large specimens of Langeites glaber are encoun

tered from a mari sam ple (collected by Anders Martinsson)

at Holmhallar l .

Ecological notes The distribution ofjawed polychaetes on Gotland seems to

a large degree to be related to sediment types. Sediments

on Gotland are of ten grouped into marIstone and lime-

Fig. 14. Hypothetic phylogenetic relationships between Gotland paulinitids. See Fig. 12 for explanation. DA. Lanceolatites gracilis, left and right MI and MIl, Nygårdsbackprofilen l , Lower Visby Beds, unit e, sample 79-42LJ. DB. L. gracilis, left and right MI and MIl, Ajmunde l , Hemse Beds, Hemse Mari, NW part, sample 71-40LJ. DC. L. gracilis var. visby, left and right MI and MIl, Vattenfallsprofilen l , Lower Visby Beds, unit e r? ) , sample 76-6LJ. DO. L. gracilisvar. visby, left and right MI and MIl, Fjale 3, Klin teberg Beds, unit e , sample 77-7CB. DE. L. sp. A, left and right MI, Strands l , Hamra Beds, unit b , sample 76-19CB. O F. Hindenites gladiatus, left and right MI and MIl, Vattenfallsprofilen l, Hogklint Beds, unit d, Valdaria testudo bed, sample from the Swedish Museum of Naturai

History (RM) . O G. H. gladiatus, left and right MI, Gandarve l , Halla Beds, sample 7l-81LJ. O H . H. gladiatus, drawn from the type speeimen (right MI) and the left MI from the same sample (sample 0.308, cf. Kielan:Jaworowska 1 966) , locality and stratigraphic position unknown, probably late Wenlockian. DI . H. angustus, left and

right MI and MIl, Vattenfallsprofilen l , Hogklint Beds, unit d, Hernnannina bed, sample RM. DJ. H. naerensis, left and right MI and MIl, Narshamn 2 , Burgsvik Beds, lower part ( ? ) , sample 83-l 2LJ. DK. Kettnerites (K) versamlis form A, left and right MI and MIl, Buske l, Lower Visby Beds, unit e, sample 79-40LJ. DL. K (K) versamlis form A, left and right MI , Håftingsklint l , Upper Visby Beds, sample 76-1 0CB. DM. K (K) versamlis form C, left and right MI and MIl, Buske l, Lower Visby Beds, unit e , sample 79-40LJ. ON. K (K) versamlis form B, left and right MI, Håftingsklint l , Upper Visby Beds, sample 76-l 0CB. D O . Gotlandites slitensis juvenile specimen, left and right MI and MIl, Ajstudden l , Slite Beds, unit Pentamerus gothlandicus, sample 83-7CB. DP. G. stitensis adult speeimen, left and right MI, Vallstena 2, Slite Beds, unit Pentamerus gothlandicus or slightly older, sample 77-2CB; left and right MIl, Vike 2 , Slite Beds, unit Pentamerus gothlandicus or slightly older, sample 83-4CB. D Q. Langeites glaber, left MI, Stora Kruse l, Hemse Beds, Hemse Mari, SE part, sample 82-29LJ; right MI, Vaktård 4, Hemse Beds, Hemse Mari, SE part, sample 81-35LJ. D R. L. glaber, left and right MI and MIl, Holmhallar l , Sundre Beds, middle upper part, sample MS907AM. O S. L. glaber drawn from the type specimen (right MI) and left MI from the same sample of unknown age (sample 0.466, Kielan:Jaworowska 1966) .


stone, but this subdivision is toa imprecise to give any

deeper knowledge of the distribution of the polychaetes.

Without a study (including thin-sections) of the sedimen

tology, the ecological discussion (which is based, moreover,

on only about one third of the polychaete fauna) , can be

only of a preliminary nature .

On Gotland, two to four paulinitid species is the normal

diversity found in a sample yielding polychaete jaws. A

low-diversity fauna, dominated by small specimens, and

often coupled with a fairly low abundance, seems to indi

cate fairly deep water. Deeper water is usually inferred

sedimentologically by a high con tent of argillaceous mate

rial , but the latter is only an indication of low water energy.

Faunas with a high diversity probably inhabited bottoms

with low water energy, i .e . a marly substrate but not from

the very deepest waters. Thus, lagoons and other restricted

environments probably offered the most varied and opti

mal conditions for the paulinitid polychaetes.

Severai samples from the limes tone are as of, for in

stance, the Slite Beds, Klinteberg Beds and Hemse Beds

have a very low abundance of polychaete jaws (Figs. 2-4 for

the abundance and the stratigraphical distribution; Figs .

5-1 1 for the geographical distribution) . This is probably

partly due to unfavourable preservational conditions, but

the faunal diversity was probably fairly Iow as well.

Most of the paulinitid species on Gotland show a prefer

ence for particular habitats. The following descriptions

should be seen as sketches, and are not inrended to cover

the total variation of the fauna, only to indicate the trends.

• Gotlandites slitensis, stenotopic with a short range, found in mari, fairly deep water.

• Hindenites gladiatus, H. angustus, and H. naerensis, ranges of moderate length except the latter species which is found on ly at a few localities, found in shallow-water sediments, in the vicinity of reefs but not within them.

• Kettnerites (K.) abraham, probably eurytopic, with a short range, found in mari deposited in deep water.

• Kettnerites (K.) jacom, more or less stenotopic with a short range, argillaceous limestone to marI.

• Kettnerites (K ) bankvaetensis, with few exceptions found in argillaceous limes tones on the eastern side of the island.

• Kettnerites (Kl burgensis, stenotopic with a short range, found in mari deposited in both deep and fairly shallow water.

• Kettnerites (K ) huberti, eurytopic with a long range, found in mari deposited in deep water, and argillaceous limestone but not 'well washed limestone ' .

• Kettnerites (K) martinssonii, eurytopic with a long range, commonly found in most types of sediment except in maris deposited in deep waters.

• Kettnerites (K ) polonensis, long range, a typical limestone-loving form but also common in the Mulde Beds (marIstone) . Taxon typical in sediments deposited in shallow waters.

• Kettnerites (K) versamiis, stenotopic with a short range, found in mari deposited in deep water.

• Kettnerites (A.) sisyphi sisyphi, not very shallow or deep water.

• Kettnerites (A. ) sisyphi klasaardensis, stenotopic with a short range, mari deposited in deep water.

26 Claes F. Bergman

• Kettnerites (A.) sisyphi var. valle, short range, typical populations

found in mari deposited in deep water.

• Kettnerites (A. ) microdentatus, long range, usually found in mari bm not in the deepest deposited sediment.

• Kettnerites (A. ) siaelsoeensis, stenotopic (? ) very short

• Lanceolatites gracilis, long range, normally found in mari deposited at moderate depth.

• Langeites glaber, range of moderate length, a very rare speeies found only in marI.

Comparison with paulinitid faunas from other are as

Due to the scarcity of reports on the jawed polychaetes and

the confused taxonomy, the regional distribution ofspecies

is not known in any great detail. The question whether the

paulinitid species were widespread or showed provincial

ism has so far been impossible to discuss since, at least, most

of the Baltic species have been treated as one species.

However, Taugourdeau (e.g. 1 976) has tried to correlate

Silurian and Devonian form taxa between North America,

France and the Sahara. In my opinion it is dangerous to

attempt comparisons based on badly preserved or partly

buried elements, or with only one or a few elements of each

type and taxon, or a combination of these features. The

identification of biological species based on well-preserved

elements of the four main types is the key to polychaete

taxonomy, and thereby to reliable regional comparisons.

Sweden. - Only a few reports have been published on fossil

jawed polychaetes from Sweden, and no paper has dealt

with paulinitids outside the Silurian strata of Gotland. The

Silurian fauna from Bjarsjolagård, south Sweden (Scania) ,

is similar to that of Gotland but the morphology of the jaws

is slightly different (courtesy of Fredrik Jerre , 1987) .

The Baltic area. - A large number of polychaete jaws and apparatuses have been described by Polish students. Most of the material derives from erratic boulders of Baltic ori

gin . However, the exact stratigraphic and geographie prov

enance of these boulders is not known and will probably

never be possible to trace. The three species Hindenites

gladiatus, Kettnerites (K.) polonensis, and Langeites glaber, described by Kielan:Jaworowska ( 1 966) from erratic boulders,

have all been found on Gotland. Her H. gladiatus resembles

the Early to Middle Wenlock representatives of the species

on Gotland. K. (K.) polonensis is very difficult to date due to

the minute size of the type specimens; an Early Ludlow age

is probable. Langeites glaber is very rare , both on Gotland

and in the erratic boulders dissolved in Warsaw; the erratic

material is transitory between the Gotland Sundre and

Hemse material . The age of the erratic Langeites glaber

fauna could be Late Ludlow. Beside these taxa, others,

which have been described in this monograph, are present

in the samples (e .g. samples described by Kielan:Jawor

owska 1966) in Warsaw. For example, in boulder 1 87/32

jaws conspecific with Kettnerites (A. ) sisyphi var. valle and in

boulder 1 5 1 /4 jaws similar to Kettnerites ( K. ) abraham isaac


are found. The small number of 'marl loving' taxa found in

erratic boulders is notable .

It is not improbable that at least some of the boulders

found in Poland derived from, or close to , the Gotland

area. Thus, a close resemblance of the faunas can be ex

pected and will not increase our knowledge about the

faunal distribution to any great extent.

Poland. - Polychaete jaws and apparatuses have also been

described from borings in Silurian rocks in Poland, e.g. a

paulinitid from the Mielnik boring (Szaniawski 1970) . The

paulinitid specimen described as Paulinites polonensis

Kielan:Jaworowska is probably conspecific with Kettnerites

(K.) huberti from Gotland.

Great Britain. - The jawed polychaete fauna from the Lud

low strata of the Welsh Borderland show a close similarity

to the Gotland fauna. In Allison Brook's unpublished col

lections from this area I have seen some species which are

also present on Gotland, e .g. Kettnerites ( K. ) huberti and

Lanceolatites gracilis (courtesy of A. Brooks) . The low diver

sity of the assemblages reported by Aldridge et al. ( 1 979, p.

437) from the Whitcliffian of the British Isles is in agree

ment with the fauna of Gotland. I do not know whether this

is caused by the local environment or whether it is due to a

global impoverishment of the polychaete fauna.

Northem Siberia. - Mannil & Zaslavskaya ( 1 985a) have de

scribed Silurian polychaete faun as from northern Siberia,

including the island Severnaya Zemlya in the Arctic Sea.

The L1andovery right MI (738/51 ) from the Waterfall sec

tion on the island of Severnaya Zemlya is very similar to the

corresponding element of Kettnerites (K. ) abraham abraham

in the L1andovery strata of Gotland. The left MI ( 738/50) ,

illustrated together with 738/5 1 is less characteristic, and

thus less easy to identify, but could possibly be the counter

part of the right MI. The two Wenlock MI jaws (Ieft MI

738/54 and right MI 738/55) from the Srednij sequence of

the same island seem to belong to different species, but the

inferior preservation and the angle at which the photo

graph was taken make them difficult to assess. However, l

believe the right MI to be conspecific with Kettnerites (A . ) sisyphi. The left MIl (738/56) from the Wenlock Srednij sequence is very similar to the corresponding element of

Kettnerites (K.) polonensis from Gotland. The right MIl

( 738/57) is older and from an other locality. They probably

do not belong to the same species.

The fauna described by Mannil & Zaslavskaya is very

similar to the fauna from Gotland, but identifications based

on single specimens are normally not enough for reliable

conclusions.

Bohemia. - The type material of Kettnerites kosoviensis Zebera

1935 consists of very large jaws, flattened and in part

crushed, partly buried in the sediment. It will not be possi

ble to dissolve the matrix to free the jaws without breaking

them. The jaws show a slight resemblance to Kettnerites ( K. ) martinssonii (described by Martinsson 1 960 as Paulinites

sp . ) .

}Tance and Sahara. - Taugourdeau (e .g . , 1 968, 1 976) has

reported paulinitid jaws from the Silurian , Devonian and


Carboniferous of France and the Sahara. He has also iden

tified them as conspecific with North American scoleco

donts.

North Amenca. - The North American polychaete fauna,

from the Ordovician to the Devonian, is rich and varied.

Eller ( 1934a, 1 934b, 1 936, 1938, 1 940, 1941 , 1942, 1945,

1955, 1 963a, b, 1964, and 1967) , Stauffer ( 1 939) , Sylvester

( 1 959) and Boyer ( 1 975) have described a large number of

elements including apparatuses belonging to paulinitids.

However, only few of the species could be placed within the

'Baltic genera' and no identical species seems to have been

described, although a jaw from the Silurian of Wisconsin

shown to me by Jeffrey J. Kuglitsch is very similar to Kettner

ites ( K. ) martinssoni (Fig. 40B) from the Hamra Beds. Walli

ser ( 1 960) described paulinitid elements from the Cana

dian Arctic Archipelago. Some of these jaws show a very

slight similarity to species from Gotland (e .g. Idraites sp.

and Polychaetaspis ? kozlowskii described by Walliser 1960) .

Material and methods

Localities and samples. - The locality name and geographical

situation, stratigraphical level, sample number, sample

level and recorded scolecodont fauna for each sample is

given in the Appendix. New localities are described in

accordance with Laufeld 1974b and localities described

earlier, with reference to the original description (s) .

In the field, each sample was given a unique sample code

in the form initiated by Laufeld & Jeppsson 1 976, e.g.

G75-7CB. G stands for Gotland, but is omitted here since

all samples (with sample number) discussed in this mono

graph derive from Gotland; 75 stands for the year of collec

tion; 7 for the seventh sample of that year from Gotland;

and the last two letters stand for the initials of the collector,

in this case Claes Bergman . The collectors of other samples

used in this publication are : Lennart Jeppsson (LJ) , Doris

Fredholm (DF) , Sven Laufeld (SL) , Sven Stridsberg (SS) . I

have also had access to a num ber of large scolecodonts

picked out by Anders Martinsson and Kent Larsson from

their ostracode and tentaculitid mari samples. The sample

code of the museum material is abbreviated as follows: the Swedish Museum of Natural History, Stockholm as SMNH,

and the Geological Survey of Sweden as SGU.

All illustrated specimens not already belonging to any

museum collection are deposited in the type collection of

the Department of Historical Geology and Palaeontology,

University of Lund, under the designation LO + a number.

Collecting. - Standard micropalaeontological and strati

graphical field methods have been used. In order to get a

proper view of the geographical and stratigraphical distri

bution, collecting was not restricted to highly productive

lithologies, although during the later years of field work, I

concentrated to lithologies that might be expected to yield

a good fauna. Rock surfaces exposed to prolonged weath

ering in recent time were avoided, since an oxidizing envi

ronment is harmful to the preservation of anne lid jaws.

The positions of the samples within each section was mea-


sured relative to suitable reference points or leve I s , such as

a bentonite layer, the base of a reef, or an artificial marker

of more permanent type. In order to facilitate fu ture col

lection at any sampled level, severai of the localities were

photographed with a polaroid camera and the sample

levels marked on the print while still in the field. (The

prints have been deposited at the Department of Historical

Geology and Palaeontology, University of Lund. ) The sam

pling interval within a section depends on the lithology.

Normally, at least two samples were collected from a local

ity, except for small exposures with uniformly developed

sediment. Sample weight varies from 3 to more than 50 kg.

To start with , I collected samples of 2-4 kg depending on

how difficult it was to collect. In later years normally about

8 to 10 kg of rock was sampled to get a better view of the

fauna. It is now evident that samples of at least 50 kg will be

necessary when dealing with the jawed polychaete fauna in

many limestone areas on Gotland.

As a standard, in a section dominated by marly layers, the

collected samples were taken from levels with the highest

calcium carbonate content. In the limestone beds, most of

the samples were taken from fine-grained sediment.

Laboratory technique. - The methods used in the laboratory

have been improved considerably over the years, chiefly by

Lennart Jeppsson (Jeppsson et al. 1 985; Jeppsson & Fred

holm 1987; Jeppsson 1 987; Barnes et al. 1 987) . Methods

invented elsewhere , published or available through oral or

written communications, have also been adopted. Most of

these changes do not effect the composition of the recov

ered scolecodont fauna, except that the increased effi

ciency has permitted larger samples to be processed. The

increased num ber of methods have also permitted a wider

selection of localities to be sampled.

The laboratory method now used is in brief as follows:

First a slab, a pilot test, of at least 0.5 kg but not less than

about one tenth of the total sample weight is cleaned

mechanically with a wire-brush . If lichens, etc . , need to be

removed, the sample is etched in hydrochloric or acetic

acid for a short time. The remaining part of the sample is

stored. The uncrushed slab is dissolved in buffered 7%

acetic acid (Jeppsson e t al. 1 985) and the insoluble residue

is washed through 1 .0 mm and 63 11m screens. All fractions (except those below 63 11m) are stored for future reference. The residues of very marly samples which do not fully

disintegrate or are difficult to rinse free of clay, are heated

to around 50°C for about 24 hours or until completely dry. Then the residue is immediately soaked in petroleum-ether

for some hours, the excess fluid is poured off, but saved,

and hot water poured over the sample . The mari starts to

disintegrate within seconds and normally the sample is

completely disintegrated after a few minutes. After sifting

and rinsing in gen tly running water the sam ple is dried and

the fossils picked out. Samples yielding abundant con

odonts or fish scales are normally density separated and/ or

separated in a Frantz Isodynamic Magnetic Separator

model L-l .

Most sample residues derive from Lennart Jeppsson 's

collections. The majority have been separated by heavy

liquid and/ or magnetic separation. Scolecodonts have nor-

28 Claes F. Bergman

mally been extracted from the light fraction, though most

heavy fractions have be en picked as well, chiefly in connec

tion with the extraction of conodonts, phosphatic brachio

pods and fish scales. The yield from the heavy fraction is

very Iow as a rule, but exceptions do occur. I have so far

avoided concentrating the scolecodonts from my own sam

ples with heavy liquid, since the first maxillaes (MI) of the

investigated taxa are hollow and very elongated with a

strongly enclosed myocoele opening that may trap sedi

ment. This would bias the result of the density separation

as well as the magnetic separation . Experiments with mag

netic separation of scolecodonts have so far been without

success. An alternative method of con cen trating the scolec

odonts is to dissolve the sample in hydrochloric or hydro

fluoric acid. This method has the disadvantages of being

poisonous and destructive to phosphatic and siliceous ma

terial . Therefore , it is feasible only if the sample has first

been broken down by buffered acetic acid and/or petro

leum-ether and the phosphatic material removed by, for

instance, heavy liquid separation. This treatment of sam

ples will probably be routine when very large samples are to

be dealt with in the future .

The dried residue is spread out on a metal picking tray

with a grid system on a white background (available from

Fema-Salzgitter as 'Hand-picking scales made of brass,

Punched ' . Address: Fa. Rudolf Stratmann, 3327 Salzgitter,

Friedrich Ebert Str. 53, West Germany) . The annelid jaws

and other fossils are picked out from the residue by means

of a single hair which has been charged electrostatically,

and dropped through one of the hoies into a microfossil

slide below the tray (Barnes eL al. 1 985) . The microfossil

slide has a light coloured rectangular bottom, with dark

lines in a grid system on it. The bottom of the slide is lightly

pre-coated with a water-soluble glue based on gum arab

icum Ueppsson 1 974, p. 5) . The sorting of the annelidjaws

is conveniently done with a fine wet sable brush (nr. 00) ,

and the jaws are at the same time glued in place.

In order to facilitate comparison the jaws are arranged

with each species in a horizontal row and homologous

elements in vertical columns. Thus, the MI 's are placed to the far left of the slide, the MII ' s to the right of the MI ' s of the corresponding species etc. A large quantity of jaws will

of course violate this organization of the slide since some areas will be overcrowded, especially those intended for

MI ' s and MII 's . The jaws are normally placed with the

dorsal side up, i .e . the myocoele opening facing down. This

organization reveals the large st number of specific charac

ters and will thus facilitate comparison between closely

related taxa, since homologous jaws are similarly oriented

and placed close to each other in the same column.

I have also worked with wet sample methods as described

by Kielan- Jaworowska ( 1 966, p. 15 ) , with the intention to

find scolecodont apparatuses. After more or less fruitless

experiments I concluded that apparatuses of the genus

KettneriLes and related genera must be fairly rare . Most of

the partial apparatuses from Gotland studied were found in

dried sample residues, because the majority of the sample

residues have been dried. Apparatuses should be more

common in wet samples, however, since drying will destroy


any pellicle surrounding the apparatuses (Kielanjawor

owska 1 966, p. 1 1 ) of most, if not all, of the taxa. The

majority of apparatuses found were more or less com

pressed and not in a good state of preservation.

The jaws forming an apparatus partly cover each other,

and are of ten more or less flattened and distorted, making

identification even more difficult than if single , well pre

served specimens are used. The higher abundance of appa

ratuses expected in wet fractions is likely to be ascribed to

small juvenile forms. I have not made much further effort

in collecting apparatuses since most of the information

they provide is also obtainable from isolated jaws.

The jaws may be bleached and become translucent by the

use of an oxidation medium, e.g. sodium hypochlorite

(Tasch & Shaffer 1961 , p. 369) or hydrochloric acid and

potassium chlorate (Kielanjaworowska 1 966, p. 1 5 ) . The

bleaching will facilitate the study of inner structures while

at the same time rendering the jaws fragile . I have bleached

jaws of different sizes but were unable to bleach large , thick

jaws to translucency - they become yellow but remain

opaque.

Illustration of speeimens

Scanning elecLran microscopy and photography. - A large mate

rial ofwell preserved elements, representing different pop

ulations from various stratigraphical levels was selected for

scanning electron microscope (SEM) studies. The ele

ments were mounted on specimen stubs with double-adhe

sive tape. In order to facilitate the orientation of an ele

men t on the stub, a small drop of water was added to the

surface of the tape and the specimen was placed in the

water. Thus, the element could easily be moved, but once

outside the drop or when the water evaporated it was stuck

to the tape. The specimens were oriented with the anterior

part towards the centre of the stub, if possible in the same

'resting position ' as on the slide . Thus, the ' light' will

appear from the same angle on all scanning micrographs, and the micrographs are taken from the same angle at which the elements are normally studied, when using a

normal stereoscopic microscope . I have not used the same specimen for the ventraI and the dorsal view, since the risk

of fracturing the specimens is very great when loosening

them from the tape. The specimens were coated with an all oy of gold and palladium in the proportions 60:40. Ini

tially, the hollow jaws tended to implode when air was let

into the vacuum-chamber of the coating device. The very

imprecise valve construction made it impossible to let in

the air more gradually with slow compression. This obstacle

was overcome by some ingenious work by Sven Stridsberg.

The vacuum pump was allowed to work and the low pres

sure could then be decreased slowly with the help of the

argon leak needle valve. After filling the chamber slowly

with argon gas for a few minutes, (easily controlled on the

torrmeter) the pump was stopped and after a light touch

on the air valve the lid could easily be opened.

The SEM work was carried out using a Leitz AMR 1 600T.

Most specimens were photographed without tilt. A pan

chromatic roll film, Agfapan 25 professional, was used. This


24x36 mm fine-grain film will reproduce any fine detail on

the 1 05x105 mm SEM screen, since only about two thirds

of the screen heigh t but all the width is used. This will not

affect the total capacity since most objects investigated here

are elongated. Further, no information is being lost using

this 35 mm film compared to 1 20 films, since the film

resolution gives about 50% overcapacity.

Agfa Rapidoprint paper was used for the prints. One

standard of magnification was selected, to facilitate com

parison. However, for practical reasons some other magni

fications were also used. The background of the photo

graphic figures was blocked with masks cut out from a

semitransparent plastic film (Dlano Amber Clear; Amber

lith; H6W 2655; cf. Bengtson 1 986) . Due to the small detaiIs

(e .g. denticles) the innermost part were in some cases

blackened with black ink.

Most of the drawings were made using a Nikon SMZ-10

stereoscopic microscope equipped with a drawing device .

Terminology

For ease of understanding, I have kept the num ber of

technical terms as Iow as possible. The terminology is to a

large extent based on Kielan:Jaworowska ( 1 966) and on the

glossary compiled and defined by Jansonius & Craigs

( 1 971 ) , Taugordeau ( 1 978) and Wolf ( 1 980) . The jaws in

dorsal position, named maxillary pieces (M) , are normally

numbered from posterior to anterior with Roman numer

als (MI-MVI) . The following glossary of descriptive terms

is intended for paulinitidjaws (Fig. 15 ) .

Basal angle. - The angle between the long axis of the undenticulated ridge and the outer margin of the basal portion of MI.

Basal furrow. - A fairly short furrow in the posterior part (basal portion) on the dorsal, outer side of MI, of ten parallei to the undenticulated ridge .

Basal plate. - A small to medium-sized right-hand jaw, closely fitted into a posterior bight or concavity of Mlr (�right MI) ; in some

speeies the basal plate is paired, the left-hand element being called the laeobasal plate .

Basal portion. - The posterior part of MI, which is widened compared with the denticulated anterior part of the jaw. The basal portion includes the inner wing, the undenticulated ridge, the basal furrow, the flange and sometimes a spur on the posterior outer part.

Bight. - A concavity in the margin of the outer and/ or posterior face of a jaw, open to the posterior, especially in the right MI, where it provides a space into which the basal plate can fit; also used for ramal arch.

Bight angle. - The angle between a line tangential to the outer margin of the shank and: ( 1 ) a line tangential to the inner margin of the ramus (as in MIl) ; (2 ) a line tangential to the posterior

margin of the flange (as in MI) .

Garner5. - Paired elements in the apparatus, situated behind the forceps and serving as a support for the posterior ends of the forceps or MI's.

Gusp. - The largest denticle, of ten the anterior one, in a series of denticles, esp. on the basal plate (if denticulated) , MIl, MIll, MIV. On the MIl's, the cusp may be preceded by one, two or very rarely three pre-cuspidal denticles. On some MIl's there are two anterior denticles of equal size forming a double cusp.


Guttingedge. - A small ridge, along the base to the apex on opposite sides of the denticle, oriented along the extension of the jaw.

Oeppsson 1 979, in conodont terminology. )

Dentary. - Series of denticles along the inner (dorsal) margin; according to the density of denticles the dentary can be den sespaced, normal-spaced, wide-spaced and paucidentate ; in some forms part or all of the dentary is edentulate (�dentary without denticles) , in which case the dentary is equal to the inner (dorsal) margin. Sometimes it is possible to subdivide the dentary into two parts, intermediate and posterior (�post-cuspidal denticles) .

Denticulated ridge. - A ridge with denticles, here used in a restricted sense, on MI along the inn er margin . The posterior part of the denticulated ridge may be without denticles. The term edentulate Oansonius & Craig 1971 ) refers to such an undenticulated part of the ridge.

Depth. - The term is defined by Stauffer 1933. I have used the term width instead.

Double cusp. - A pair of cusps on MIl, often with parts of equal size.

Falx. - The extension of the outer anterior part of the jaw, esp. in MI, of ten sickle-shaped, forming a hook or a fang.

Falcal arch. - The concave part of the anterior, inner margin, which may be denticulated or undenticulated Oansonius & Craig 1971 , emended herein. )

Fang. - The pointed, recurved termination of MI , less developed than a hook.

Flange. - A wing-like extension on the dorsal side of the posterior, outer part of the right MI. It is of ten more or less elevated, and if present the basal plate is fused to it. The term was used by Sylvester ( 1 959) to include only the ligament rim.

Head. - The anterior, wider part of the carrier.

Hook. - The very large, recurved anterior termination of MI is denominated hook. The less developed anterior termination is named fang.

Inner margin. - Ajaw seen in dorsal view has its inner margin along the side which is oriented facing the central axis of a paulinitidjaw apparatus, along the dentition and inner wing of the MI and MIl elements.

Inner wing. - The posterior inner margin may be developed as a longitudinally elongated and laterally extended or downfolded and extended area.

Intermediate dentary. - A group of small, of ten equally sized denticles, usually two to eight in number, on the paulinitids. They are immediately posterior to the cusp of the left MIl, and are less well-developed on the right MIl.

Length of the element. - The largest dimension of a jaw paralleI to its long axis (i .e . approximately parallei to the dentary) between the most posterior and the most anterior points.

Ligament rim. - A narrow structure (of ten a small ridge, groove or a combination of these) surrounding the myocoele opening along the anterior, the inner and the outer sides. Sylvester ( 1 959) used the term jlange for this structure.

Myocoele. - The space inside (pulpal or muscle cavity) , and more or less enclosed by the jaw, extending to the tip of the fang, hook or cusp. (Partly from Kielan:Jaworowska 1 961 ) .

Myocoele apening. - The outline of the ventrai margins of the jaw enclosing the myocoele.

Duter wing. - The posterior outer margin of the MI may be peveloped as a longitudinally elongated and laterally slightly extended area which is part of the margin of the myocoele opening.

30 Claes F. Bergman

I M I

anterior _ I a l x

e;..-- ' I N � \a l C a i a r c h I i n n e r m a r g i n

I n n e r

rMI

p o s t e r i o r

b a s a l l u r r o w densely spaced

c o m b - s h a p e d d e n t a r y

s m a l l d e n t i c l e s

I t>' \ r. i I

normally spaced

d e n t a r y

c u t t i n g e d g e

d e n t a r y

s p u r

f l a n g e

w i d e l y s p a c e d

d e n t a r y

n o r m a l - s i z e d d e n t i c l e s

FOSSILS AND STRATA 2 5 ( 1 989)

f a n g

l e n g t h

s p u r

poster ior m a r g i n � p a u c i d e n t a t e d e n t a r y

l a r g e d e n t i c l e s

d o u b l e c u s p a n t e r i o r p r e - c u s p i d a l d e n t i c l e b i g h t a n g l e

ramus

bight

� d e n t i c u l a t e d

r i d g e

i n n e r w i n g m y o c o e l e o p e n m g

p o s t e r i o r

+-- w i d t h ---+

r a m u s

c u s p ---1-T

_ w i d t h �

l e n g t h

l e n g t h

u n d e n t i c u l a t e d

r i d g e

b a s a l a n g l e

o u t e r m a r g i n

. '\"---.. b a s a l v p o r t i o n

b a s a l p l a t e

Fig. 15. Schematie drawings of paulinitidjaw elements to illustrate the morphological terms used. All are elements seen from the dorsal side , except the right figure of the left Mil element (ventraI view to show myocoele opening) .

Paucidmtate dmtary. - With few and widely spaced denticles (see dmtary) .

Post-cuspidal dmticles (Post-cuspidal dmtary) . - The den ticles on the posterior side of the cusp and posterior to the intermediate dentary. The term posterior dentary is also sometimes used (see dentary) .

Pre-cuspidal dmticle (pre-cusp) . - Any denticle an terior to the cusp or a small denticle on the anterior side of the cusp (see dmtary) .

Ra.mus. - Any arm-like lateral extension of the face of a jaw projecting from the over-all outline, usually pointing posteriorly, esp. in MIl, MIll, and MId (;right MI) ; in some forms pointing anteriorly, especially the basal plate . (Jansonius & Craig 1971 . )

Scalecodont. - Any jaw piece of a polychaetous anne lid; originally intended only for isolated fossil elements. (Croneis & Scott 1933.)

Skaft. - The posterior extension of a carrier or a mandible.

Skank. - The undenticulated posterior extension on the inner side of ajaw. The most posterior part of the shank can have a blunt or a sharp termination.

Slantingdmticles. - Denticles pointing in the posterior direction.

Spur. - A short, ridge-like process which is a prolongation of the thickened margin of the myocoele opening on MI. The spur can of ten be seen in dorsal view on the outer margin on the anterior part of the basal portion.

Undmticulated ridge. - The ridge on the basal portion of MI, extending as a continuation of the denticulated inn er margin (also referred to as edentulate by other authors) .

Vmtral side. - The side of the jaw to which the myocoele is open.

Widtk. - The largest dimension of a jaw perpendicular to its long axis and on a plane paralleI to the myocoele opening. The width of the MI is measured anterior to the basal portion.


Structure , composition, and preservation of polychaete j aws

Chemical composition. - It has of ten been claimed that the

jaws of polychaetes are composed of chitinous material

(e .g. Schwab 1 966, p. 416) . This is, however, not the case

according to Voss-Foucart et al. ( 1 973) , who demonstrated

that Recent polychaete jaws are composed of scleroprotein.

Olive ( 1 980, pp. 572, 580) has published a thorough review

of the much-varied composition of the jaw material in

different taxa. The jaw material includes tanned protein,

aromatic amino acids, glycerine, hystidine and metals such

as iron, zinc and copper. Recent eunicids have a high

concentration of calcium carbonate , while the phyllo

docids have less. Colbath ( 1 986) suggests that the heavily

sclerotized jaws with little mineral reinforcement have a

better prognosis for fossilization . Some of the fossil eunicid

jaws might have had the same reinforcement of calcium

carbonate as the Recent forms have (Colbath & Larson

1 980, p. 485) , though Boyer ( 1981 ) sugge sted that the

calcium carbonate might have been replaced in the jaw post

morfem. Some fossil eunicid jaws also include fluorapatite,

but whether this is a reinforcement or of secondary origin

is an open question according to Schwab ( 1966, p. 42 1 ) .

The Recent jaws studied lack fluorapatite (Schwab 1 966, p.

421 ) , although they are of different phylogenetic lineages

than the fossil jaws investigated, and therefore give little

information about the origin of the fluorapatite .

Preservation. - It has also been claimed that scolecodonts are

very resistant to acids, oxidation, thermal alteration and to

changes due to the recrystallization of the carbonate ma

trix (Jansonius & Craig 1971 , p. 251 ) . However, Kozur

( 1 972, p. 753) states that the resistance towards oxidation

and matrix recrystallization is very poor. From my material

of the Lower Palaeozoic of Sweden it seems quite clear that

scolecodonts are not common in any highly oxidized (red)

sediment. Szaniawski ( 1 974) and Colbath ( 1 986) demon

strate that there is a considerable variation in the resistance

to chemical oxidation among the jaws of different poly

chaete families.

Thejaws have almost the same colour variation as amber:

from brownish black through reddish brown to yellowish brown. The colour varies between some taxa but is consistent within the species apart from the sm alle st, of ten translucent, jaws. There is no sign of the jaws having been

affected by high temperature . Dorning ( 1 984) noted the

possibility of using different types of fossils, including polychaete jaws, to enhance the precision of thermal metamor

phosis indexes such as those based on conodonts (Epstein

et al. 1 977) . Laufeld ( 1974a, p. 1 28) concluded that the

Gotland chitinozoans had never been subjected to temper

atures exceeding 1 00°C, andJeppsson ( 1 983, p. 1 22) found

no sign of thermal alteration in the conodonts. This im

plies a maximum temperature of less than 80°C.

The preservation of scolecodonts from the Silurian of

Gotland varies from oxidized, flattened, more or less un

identifiable fragments to beautiful three-dimensionally

preserved elements without any sign of abrasion or com

paction. Large elements, more than 1-2 mm in length,

Silurian paulinitid polychaetes from Gotland 3 1

seem to b e more easily oxidized and brittle, and thus more

susceptible to fragmentation than smaller ones. In most

soft mari samples the jaws are more or less flattened,

whereas in the more competent calcium carbonate-rich

layers, the jaws are normally preserved in full relief. Many

limestones consist of well washed, more or less worn skeIe

tal debris, e .g . the lower part of the Slite Beds and most of

the Klinteberg Beds. The scolecodonts are normally very

rare and fragmented in these limestones, although in some

samples very well preserved specimens are encountered. In

the oolitic beds only some undeterminable fragments of

scolecodonts have been found.

The best preserved elements are found in fine-grained

sediments with a high carbonate content. I have not noted

nor searched for any effect of sediment recrystallization on

the scolecodonts (cf. Kozur 1 972, p. 753) .

In severaI localities, soft marIstone is intercalated by

competent argillaceous limestone. In the Lower and Upper

Visby Beds, for instance, the limestone is of ten in the form

of irregular nodules or thin lenses. This ' cyclic bedding'

could be a result of a rhythmic change of the physical

conditions and/or of mass transportation of ei ther the clay

or the carbonate-rich component, but it could also be of

diagenetic origin. The present work is not intended to

interpret the sedimentology be hind the ' cyclic bedding' ,

but i n samples from the Upper Visby Beds and Hogklint

Beds (e .g . , Haftingsklint l and Vattenfallsprofilen l ) no

difference in the composition of the jawed annelid fauna is

noted between marIstone and limestone within a cycle .

Transportation of any of the sediment components is not

supported by the distribution of the scolecodonts, but

bioturbation affecting only the annelid fauna could have

occurred. In a similar type of bedding in the Hamra and

Burgsvik Beds with dominating limestone sediments, a

change of the jawed annelid fauna is noted, e .g . in Bankvat

l (Hamra Beds) , but not in Glasskar l (Burgsvik Beds) .

Because no significant difference in the annelid diversity

was found between limestones and adjacent mari beds,

samples collected in the most recent years mainly derive

from the competent limestone layers. Still, less than one

sample in ten yields an abundant and well preserved fauna.

Jeppsson ( 1 983, p. 1 22) notes the same condition in con

odonts, adding that the best conodont samples of ten yield

excellently preserved scolecodonts without any sign of deformation.

Ultrastructure. - Only a few reports on the ultrastructure of

the external and internal surfaces of annelid jaws have been published, such as Taugourdeau' s ( 1 972) study of the

ornamented cuticle of Palaeozoic polychaete jaws. This

limited interest is probably due to the usually smooth sur

face of the jaws. Light-microscopy studies by Tasch & Shaf

fer ( 1961 ) and Schwab ( 1966) on bleached jaws, were

superficial in nature . The increasing use of transmission

electron microscopy (TEM) and particularly scanning elec

tron microscopy (SEM) has advanced the study of ultra

structures: almost simultaneously, severaI papers describ

ing the ultrastructure of annelid jaws appeared, e .g.

Strauch ( 1 973) , Corradini, Russo & Serpagli ( 1974) , and

Mierzejewska & Mierzejewski ( 1 974) . Strauch ( 1 973) noted


a layer with a smooth non-porous surface covering three

different internal layers with pore-canals in different direc

tions. He suggested that pore-can als could, among other

functions, have increased the elasticity. The result was in

general confirmed by Corradini et al. ( 1 974) , who worked

with a larger material and made a more penetrating study.

They distinguished a fourth layer and also studied the

different pore-canals, attempting to interpret their func

tion . Further, they noted exceptions to the smooth dorsal

surface in some taxa, e.g. Polychaetaspis tuberculatus, which

displays striation and very small pores. Shallow furrows

were noted on the inner face of Mochtiella. Severai different

types of structures, induding pores and what looks like

small corroded pits, were recorded on the ventrai side .

Charletta & Boyer ( 1974) noted striation on dentides of

Arabellites auburnensis. The taxonomic importance of the

ultrastructure, hinted at by Corradini et al. , was demon

strated by Szaniawski & Gazdzicki ( 1 978, pp. 7-1 0) .

I have not noted any ultrastructural pattern on the

smooth dorsal surface of unaltered paulinitid jaws from

Gotland. Eroded dentides (Fig. 1 6B) show layers (Fig. 1 6A)

similar to the ones described by, e .g . , Corradini et al.

( 1 974) . However, on the ventrai surface around the myo

coele opening, pores similar to the ones illustrated by

Corradini et al. ( 1 974, p. 1 26, pl. 42) have been found (Fig.

1 6C, D, E) .

Dental formula. - Severai types of dental formula exist. The

simplest one gives the total range of the num ber of denti

des of the complete jaw. This number is usually not of

major importance. A more significant type of dental for

mula was proposed by Taugourdeau ( 1 968) and later mod

ified by Jansonius & Craig ( 1971 , pp. 261-264) and Tau

gourdeau ( 1 978, pp. 1 4-17) . Germeraad ( 1 980, p. 5) noted

that with simple or very complexly denticulated jaws it is

easier to understand the dentation from an illustration

than from a complex codification.

In my opinion, existing formulas are generally difficult

to re ad and compare , and are typographically complicated.

It is at least as simple to draw a profile of the dentition as to try to work out the form ula for a complex denticulated jaw. Dental formulas are potentially useful for computerized comparisons, but with the present Silurian material the

form ula would have to express the number, direction, size

and shape of the cusp, precusp (s) , intermediate dentides and posterior dentides, not only small-large, but also in a

more differentiated form. For the present purpose , formu-

Fig. 16. DA. Three deeply eroded denticles from the middle part of the denticulated inner margin af the left MI, Kettnerites (K) bankvaetensis, LO 5841 : 1 , Glasskar 3 , Eke Beds, 82-18CB, x 1700. DB. Deeply eroded anterior part of the denticulated inner margin, right MI, K (K) bankvaetensis, LO 5841 :2 , same sample as A, x325. DC. Close-up of the pores of the ventrai surface within the posterior part of the pulp cavity of the left MI of K (K) bankvaetensis, LO 5829:8, Bankvat l, Hamra Beds, unit b, 82-30CB, x2000. OD. ventrai view of part of the basal portian shown in C, same spe eimen as in C, x5 10 . DE. ventrai view of the auter half of the basal portion of right MI, K (K) bankvaetensis, LO 5829:7, same loeality as C,

x500.


las would be an unnecessary complication, and thus I have

not used any type of dental formula besides stating the

range of the number of denticles of the jaw.

Molting of j aws and ontogeny

Jaws may grow continuously (only possible with simpler jaw

forms, such as a conical type) , or by shedding. It is also

possible that wom jaws were shed, although this is less

probable. Continuous growth is demonstrated by growth

lines in some jaws, e .g. mandibles (Paxton 1 980) . Growth

lines have not been reported from eunicid maxillae, but

are common in those of the Phyllodocida, e .g . , Nephtyidae,

Glyceridae and Nereidae (Herpin 1 926, Schwab 1 966 and

Charletta & Boyer 1 974) .

Molting of jaws in Recent annelids has been described or

suggested in a num ber of studies (summary with references

in Kielanjaworowska 1 966, pp. 49-5 1 ) , for example that of

Herpin ( 1 926) on the recent phyllodocid Odontosyllis clena

stoma. Molting is also described from the intensively studied

Recent eunicid genus Ophryotrocha (e .g . , Bonnier 1 893;

Korschelet 1 893; Heider 1 922; Åkesson 1 973;Jumar 1 974) .

A closer study of molting and the morphological develop

ment of the jaws of Ophryotrocha maculata is given by Åkes

son ( 1 973) , who studied a population of about 300 individ

uals for more than two years. Åkesson concludes that the

jaws are replaced at intervals, and that an ontogenetic

dimorphism occurs. Similarly, the jaw morphology of O.

puerilis changes during ontogeny (Pfannenstiel 1 977) . Pax

ton ( 1 980, p. 545) claims that among most Recent eunicids

the jaws grow continuously during life and produce jaws

without growth-lines. However, also eunicimorphjaws that

are shed lack growth-lines. Recently Colbath ( 1987a) con

cluded that the internal structure of maxilIae in eunicoid

polychaetes is inconsistent with continuous growth, and he

therefore rejects Paxton's theory.

Evidence for molting in fossil annelids is difficult to

verify, however. Kielanj aworowska ( 1 966, pp. 50-5l )

noted that there was no evidence of mol ting in fossil labido

gnath and prionognath types, but suggested that in placo

gnaths, assemblages of morphologically identical jaws arranged as a jaw-within-ajaw structure could indicate molting. Schwab ( 1 966, p. 420) also reports nested placognath jaws belonging to the genus Staurocephalites, the sec

ond jaw being arranged within the firstjaw, with the same

alignment in every case . Schwab compared this with tooth replacement in mammais, but did not entirely exclude

other explanations, such as growth abnormalities. Such

phenomena have also been noted among the same type of

jaws from the Silurian of Gotland. Mierzejewski ( 1 978a)

discussed different types of molting and evidence in sup

port of it. He further stated that any increase in the size of

eunicid jaws during ontogeny must occur by molting since

a biochemical change occurs in the jaws which excludes the

possibility of growth by accretion . Based on internal studies

of Recent material, Colbath ( 1 987a, p. 446) proposes that

palaeontologists should tentatively assurne that moulting

occurred among the jawed fossil polychaetes.


The ontogenetic development shown in Kettnerites ( K. )

martinssonii and Gotlandites slitensis i n this monograph, is

also evidence of jaw replacement, if the possibility of re

sorption and accretion of the jaw material is excluded ( see

Mierzejewski 1 978a) .

Thus, in accordance with Colbath ( 1 987a) I believe it is

reasonable to assurne that the jaws of many Lower Pal

aeozoic eunicids were shed and replaced during growth.

Size frequency classes. - The shedding of jaws could theoreti

cally be proved by the identification of different, delimited

size frequency classes, similar to the different instars, for

instance, among the ostracodes (e .g . Martinsson 1962 ) .

Olive ( 1 980) has stated the difficulty of obtaining correctly

delimited size classes of complete worms in Recent mate

rial. According to Olive (personal communication, 1 986)

only the first and the second size classes of the polychaete

group measured seem to be clearly delimited from the rest.

The larger classes tend to merge, depending on individual

differences in growth, caused by differentiated food supply

and genetic differences.

The study of growth-lines of some Recent, potentially

long-lived species within the family Nephtyidae permitted

a size-independent estimate of age (Olive 1 977) . No

growth-lines have so far be en recorded from the eunicids,

however.

A num ber of factors may affect a polymodal size-fre

quency distribution of the jaws. Three different reproduc

tive strategies of Recent polychaetes have been recognized

(e.g. Olive & Clark 1 978) . Reproduction may occur once

during a lifetime, annually, or severai times per year during

an extended breeding season. How many times did the

normal annelid replace its jaws? When did they replace

them - cyclically or whenever needed? The shedding could

be coupled to any regular phenomenon, such as lunar

cyclicity or seasonal variations, for example in the abun

dance of prey. Is it then possible to identify the different

stages, or do the stages partly overlap each other?

If shedding took place at the same periods ih life, it

would probably be possible to distinguish at least the differ

entjuvenile stages . Variable growth rates between different

populations would obscure the pattern, however, and so

would mixing of material of different stratigraphic age due

to bioturbation or other kinds of reworking, and limits to sampling resolution. Transportation is also a possibility to be bome in mind, though Schiifer ( 1 972, p. 1 76) noted

that annelid jaws from the North Sea do not seem to be

subject to any sorting by currents. However, accumulation ofjaws may be caused by organisms preying on polychaetes.

Transportation caused by density currents is likely to have

occurred in the Silurian shallow shelf in the Gotland region

(e.g. Bergman 1981a, 1 984) . Brenchley ( 1 979) concludes

that among species of Recen t Eunicidae and N ereidae the

fossilization potential of the jaws is highly variable depend

ing on species and place of deposition.

Other obstacles hindering the clear identification of size

classes could be a changing of the rhythm of shedding

during the ontogeny, or a very large num ber of molt stages .

Less probable is perhaps size-expressed polymorphism

(e.g. male-female) . In the present material no dimorphism


•

N = 1 3 4 I r Il . I • • 0.20 . 2 2 .24 . 26 .28 . 3 0 . 3 2 . 34 .36 . 3 8 .40 .42 .44 .4 6 .48 . 50 .52 . 5 4 .56 .58 .60 .62 .64 .66 .68 .7 0 .72 .74 .76 .7 8 . 8 0 .62 . 84 .86 . 88 . 90

length in m m

Fig. 17. Histogram showing the size-frequency distribution o f 1 34 right M I elements o f Kettnentes (K.) bankvaetensis from the locality Bankvåt l , Hamra Beds, unit b. The sample comes from a 30-40 mm thick bed representing a geologically short time interval. No obvious size-frequency cIasses can be observed. Studies from other be ds and localities have given the same result.

implying sex-based differences in the apparatuses has been

found. I have attempted to find polymodal size-frequency

distributions at some localities (Fig. 1 7) yielding well pre

served paulinitid faunas, but the result was inconclusive .

Olive (personal communication, 1 986) states that with Re

cent material it is virtually impossible to identify size classes

beyond the second class.

General taxonomy Morphological variation in a species exists in both time and

space . Reconstructions of speeies distribution should be

based on populations from different geographical are as

and stratigraphical leveis; scattered observations of one or

a few speeimens are of limited use . The study of popula

tions can reveal variation within groups that exchanged

genetic information and thus forrned a speeies during a

speeific time. The imprecise dating of geological samples

will give a blurred time resolution, which can increase the

actual variation but may help us to follow the speeies

through time. 'An evolutionary speeies is a lineage based

. on a sequence of populations, evolving separately from

others with its own unitary evolutionary role and

tendeneies' (Simpson 1 96 1 , p. 1 53) .

The taxa recognized in this study are by necessity based

on jaws. The jaws are only a minor part of the animal, and many changes in the genetic code could take place without affecting the jaw apparatus. Further, a modification of the

genetic co de could affect the morphology of more than one part of the animal . But even so, some changes of

connected characters would not necessarily mean that a new speeies has evolved. Species boundaries in time are

largely subjective .

Variation within populations of ten leads to genetic poly

morphism (Ford 1 965) , which in wide-spread spe eies could

increase and lead to polytypic speeies (Cain 1 963) . Study

ing Devonian conodonts of gre at variability, Murphy et al.

( 1 981 ) replaced the previously used morphologic classifica

tion with a biologic one in which each taxon included a

num ber of morphotypes. In my scolecodont material from

Gotland, the lineages are not as complex as in the above

mentioned con odont example, perhaps as a result of the

very limited geographical area sampled. Instead of intro

dueing morphs, which might be premature concerning

jawed polychaetes, I have chosen to use a concept of infor

mal varieties when dealing with the more problematic poly

chaete populations. These varieties are to be regarded as

infrasubspecific categories with no standing under the

ICZN (Article 45e, g) . Most varieties are described sepa

rately in order to faeilitate transformation to a formal spe

eies-group taxon, if found necessary.

The criteria for delimiting the species could be speeified

for each element, i .e. for the four large posteriorjaws which

I am at present using. However, not every small change of

the morphology of a jaw is used to base a new taxon on.

Thus, the number of species are held at a lower level, which

makes it easier to follow the stratigraphical, geographical,

and ecological distribution of the speeies.

Criteria for delimiting genera are of course much more

complicated since the genera involved are closely related

and therefore have the same element arrangement.

The species identified can mostly be found in severai

leve Is through the stratigraphical sequence of Gotland. If

in an evolutionary series there is a distinct change in the

morphology of the jaws (e .g . K. (A . ) sisyphi sisyphi and K. (A.) sisyphi klasaardensis) and the two forms do not exist at

the same localities, then I treat the new form as a subspe

eies. If the two forms co-occur in the same sample I place

the latter form as a variety of the first (e .g. K. (K.) polonensis

and K. ( K. ) polonensis var. sjaustre ) .

Variability of jaws and apparatuses

The laboratory technique was designed to produce large numbers of jaw elements for population analysis, rather than to bring out the rare complete jaw apparatuses. Con

sequently the apparatuses described are reconstructed

mainly on the basis of isolated jaws from a large num ber of

localities. However, some more or less complete apparatuses were used in the reconstruction work.

The taxonomical value of Recent polychaete jaws has

been under debate (e .g . Claparede 1 870, p. 24; Treadwell

1 92 1 , p. 7; Wolf 1 976, p. 55; Wolf 1 980, pp. 97-99) , proba

bly due to morphological variation within some of the

Recent polychaete species (e .g. Kielanjaworowska 1 966;

Fauchald 1 970; Jumars 1 974) . The intraspeeific variation is

expressed in the position of elements in the apparatuses as

weU as in their syrnmetry (e .g . the syrnmetrical pair of the

MIll; or the appearance of a basal plate in the left MI,

normally developed only in the right MI) . The num ber of

elements may also vary within a species. Kielanjaworowska

( 1 966, pp. 30-38) has reviewed Recent eunieid taxa exhib

iting variation in num ber of denticles, of ten not correlated

to jaw size. This com mon variability is found in Arabella

iricolor, Diopatra cuprea, Eunice jloridana, Halla parthenopeia,


and Onuphis eremita. The number of dental plates and

element shape are also subject to variation. This seems less

com mon among Recent specie,s but is reported from Ag

laundes fulgida by Fauvel ( 1 919 , 1 953) and Hartman

( 1 944) , Arabella incolor by Kielanjaworowska ( 1 966) , Ara&

eUa mulans by Crossland ( 1 924) , and Diopatra cuprea by

Kielanjaworowska ( 1 966, p. 32) . Barnes & Head ( 1 977)

note the variation in the number of paragnaths in Nereis

diversicolor.

Change in jaw shape during ontogeny is reported from

the genus Ophryolrocha (e .g . , Bonnier 1 893 on Ophryotrocha

puerilis and Åkesson 1 973 on O. maculata) . Kozlowski ( 1 956,

pp. 1 83-1 85) , in his pioneer study of fossil jawed annelids,

demonstrated the ontogenetic development of the Pal

aeozoic Polychaetaspis wyszgordensis Kozlowski. Such dimor

phism as shown by Åkesson ( 1 973) must be very difficult to

record on fossil material. However, Szaniawski & Wrona

(manuscript in press, H. Szaniawski, personal communica

tion) have been able to distinguish such an example on

fossil material . Other examples of minor ontogenetic de

velopment are also shown in the present study, e.g. in

KeUneriles (K. ) martinssonii and Gotlandites slitensis. The

change of shape occurs late in the individual growth; it

probably represents sexual maturity.

A second type of den ticle variation occurs in some of the

Gotland taxa (e .g . Kettnerites versabilis, K. (K.) martinssonii,

K. (K.) burgensis, and Lanceolatites gracilis) . Within jaws of

equal size, two different denticulation types occur, one with

thin, densely spaced denticles and one with coarse denti

cles. The num ber of denticles varies within both types but

the coarse type has fewer denticles. A similar type of varia

tion is known from other animal groups, for example

conodonts (Serpagli 1 967, p. 54 described thinly and

coarsely denticulated elements within the same genus) .

Kielanjaworowska ( 1 966, p. 38) noted that the variability

in jaw symmetry may cause taxonomic oversplitting.

Paulinitid MII jaws from Gotland have a very consistent

dentition formula (Figs. 1 2 , 13 and 1 4) , though some

variation exists. The right MIl of Kettnerites ( K. ) bankvaet

ensis has normally one small and one large pre-cusp, but in

about 2-3% of the jaws there are two (in one case even

three) small denticles and one large . A geographically and

stratigraphically dense sampling of well-preserved material

makes it possible to identify severai such cases of intraspecific variation.

Morphological features of taxonomical importance

MI and MIl elements carry severaI taxonomically signifi

cant characters, although less so in the left elements. The

left MI of two taxa may overlap morphologically, making

identification based solely on this element almost impossi

ble . This is not common in Gotland material , but it occurs,

for instance, in some samples from the Halla-Klinteberg

boundary at Gothemshammar, where Kettnerites (K.) bank

vaetensis and K. (K.) polonensis occur together. In younger

strata the morphology changes, and it becomes fairly easy

to differentiate the two taxa solely on the left MI.


Jaw size in taxa and populations is useful for identifica

tion, though being also environmentally dependent it must

be used with care .

Features that are usually specific at the species-subspe

cies leve I include :

Right MI. - Denticle number, size and form, extension of

the dentary along the inner margin, transition from the

posteriormost denticles to the undenticulated ridge , varia

tion in denticle num ber versus jaw size, shape of the inner

margin, form of the inn er wing, sharp- or blunt-ended

shank, fused or isolated basal plate, form, thickness and

elevation of the flange, presence or absence of spur.

Left Ml. - As for the right MI, except that the posterior part

of the jaw (including the lack of basal plate) displays fewer

traits of taxonomic importance.

Right and left MIl. - The right MII ' s exhibit the most impor

tant characters (Fig. 18 ) , including number, size, shape,

position, and direction of precusp (s) (0-3) and cusp (s) ( 1

o r 2) . The cusp may be slender o r swollen at the base . The

most com mon num ber of intermediate denticles is 3-5 , but

varies. The delimitation of the intermediate dentary from

the posterior dentary may sometimes be difficult. Size and

form of the ramus and the shank, including the shape of

the most posterior part of the shank are also important

features.

Other elements which I have been able to identify in the

Silurian eunicid apparatuses include an unpaired left MIll,

left and right MIV, and carriers. However, this is only rare ly

possible, because of the few morphological characters on

the jaws.

The mandibles are black, massive and of ten fragmented.

Partly due to their usually bad preservation and thereby low

number in the samples in relation to the maxillae , I have

not assigned the mandibles to taxa.

Since most of the polychaete jaws are hollow they are

of ten found more or less squeezed, thus creating identifi

cation problems. Some of the above mentioned criteria will

still be possible to use, e .g . the dentition form of the

anterior denticles of the MIl and the dentition of the MI.

However, I believe it is almost impossible to base an identi

fication on flattened material without being familiar with a well preserved similar fauna (preferably preserved in full relief) .

In the MIl, the anterior part of the dentary, and in the

MI, the dentary and the basal portion (especially the shape

of the posteriormost tip of the shank) seem to be very

persistent through time. These features might prove to be

very important in phylogenetic studies.

Scolecodonts versus apparatuses

The natural taxonomy of fossil annelids is so far mostly

based on jaw apparatuses. The scolecodont-based species

described earlier are rare ly illustrated in a way that permits

comparison and identification from the photo or drawing.

This has led severaI students to ignore these taxa without

studying the type specimens. A large number of taxonomic

36 Claes F. Bergman

K. ( K . ) p o l o n e n s i s

K . ( A. ) m i c r o d e nt a t u s

L a n g e i t e s g l a b e r

M

K. ( K o l p o l o n e n s i s

v a r . g a n d a r v e

K . ( K . ) b u r g e n s i s

K . ( K . ) b a n k v a e t e n s i s

K . ( A . ) s i s y p h i s i s y p h i

K . ( A . ) f j a e l e n s i s

K. ( A . ) rn i c ro d e n t a t u s

FOSSILS AND STRATA 25 ( 1989)

K . ( K . l m a r t i ns s o n i i

c

K . ( A . ) s i s y p h i var. val le

• K . ( A . ) s i a e l s o e e n s i s

K . ( A . ) m i c r o d e n t a t u s

L a n c e o l a t i t e s g r ac i l i s l a n c e o l a l i t e s

g r a c i l i s v a r . v i s b y


Table 1. A key to the identification of paulinitidjawed polychaetes from Gotland based on the right MIl element. An asterisk ( * ) denotes that a speeies occurs in two different groups, i . e . they have a varied dentition formula.

l . With cusp (s) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (see) 3 2. Without cusp or no pronounced cusp (s) :

A. Kettnerites (K ) polonensis var. gandarve B. Hindenites naerensis

3. Single cusp . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (see) 6 4. Double-cusp:

A. Hindenites angustus 5. Double-cusp with small denticles in between (one, or rarely two) :

A. Lanceolatites gradlis B. Lanceolatites gradlis var. visby C. Lanceolatites sp. A

6. Single cusp with large pre-cuspidal denticle (s) . . . . . . (see) 1 1 7 . Single cusp with small pre-cuspidal denticle (s) . . . . . . . (see) 9 8. Single cusp without pre-cuspidal dentary:

A. Kettnerites (K ) versabilis 9. Single cusp with one small precuspidal denticle :

A. Kettnerites (A. ) siaelsoeensis B. Kettnerites (A. ) microdentatus* C. Kettnerites (A. ) sisyphi klasaardensis D. Hindenites gladiatus E. Gotlandites slitensis (precuspidal denticle of fairly small to

moderate size) 10. Single cusp with two small precuspidal denticles:

A. Kettnerites (A. ) microdentatus* B. Kettnerites (A. ) fjaelensis C. Kettnerites (A. ) sisyphi sisyphi D. Kettnentes (A. ) sisyphi var. valle

I l . Single cusp with one large precuspidal denticle: A. Kettnerites (K ) burgensis* B. Kettnerites (K ) martinssonii

1 2 . Single cusp with two large precuspidal denticles: A. Kettnerites (K) polonensis B. Kettnerites (K ) polonensis var. sjaustre C. Kettnerites (K) sp. A D. Kettnerites (K) alYraham alYraham E. Kettnerites (K) alYraham isaac F. Kettnerites (K ) jacobi G. Kettnerites (K) huberti H. Kettnerites (K) burgensis* I. Langeites glaber J. Kettnerites (K ) bankvaetensis ( the posterior denticle of the precuspidal dentary is usually smaller than the an terior one)

Fig. 18. Guide to the identification of the paulinitid jawed polychaetes from Gotland. The key element is the right MIl, which is usually the most varied element, together with the right MI. The best features for identification down to species/ subspecies level are the number, size and arrangement of the pre-cuspidal denticles and the cusp of the right MIl. The arrows point out further details of importance for the identification, such as the convexity in the outer side of the ramus and sharp pointed shank. Two opposing arrows on opposite sides denote a narrow shank or ramus. Two parallei arrows on the posteriormost part of the shank indicate the almost parallei sides of the shank. Two triangles pointing away

from each other within the ramus denote a wide ramus. Additional characteristics in the other element-types are sometimes important for proper identification (see Tab. l and the respective descriptions) .


names based on a fragment or just a few jaw specimens,

have thus been left out of consideration.

To erect a biological taxonomy of any organism paraUel

to an existing form taxonomy based on fragments is against

the rules of the International Code of Zoological Nomen

clature ( 1 985 e.g. article 23) . Eller ( 1 964) , Tasch & Stude

( 1 965) were against the use of parataxonomy, and they

argued for a combination of the two taxonomies. Kozur

( 1 970, 1971 ) tried to combine the two different systems in

accordance with the rules, but only to the generic level. His

attempt was sketchy and was rejected as premature by

Jansonius & Craig ( 1 971 ) . Szaniawski & Wrona ( 1 973) also

referred to Kozur's taxonomical ide a as premature and

criticized severai of his interpretations.

After a review of the apparatus-based genera, Edgar

( 1984, pp. 256, 260) noted that posterior elements (e .g .

MI) in the Palaeozoic apparatuses are identifiable at the

genus level, both isolated and as part of an apparatus. In

my opinion an identification at the specific level using the

MI is also possible.

Descriptions and reconstructions ofjaw apparatuses. - The first

described apparatuses were found on bedding-planes (e .g . ,

Massalongo 1 855; Hinde 1 896; Zebera -l935; Lange 1 947;

Martinsson 1960) . The jaws are either fused together or

lying on the bedding plane, like a cluster, more or less

isolated. These finds are due to chanee and/or a careful

study of the sediment surface . When the etching method

of searching for complete apparatuses was introduced

(Kozlowski 1 956; Kielan:Jaworowska 1 966) it was possible

to make more than just unique finds of apparatuses.

This technique avoids rinsing of the acid resistant resi

due through a sieve : instead the dispersed clay particles are

poured out, leaving the larger particles, including the fos

sils, on the bottom of the vessel; the residue is studied when

still under water (Kielan:J aworoska 1 966, p. 15 ) .

Study of complete apparatuses makes it possible to ap

preciate the size relations between the elements in an

apparatus, but the identification of single elements within

the apparatus might be more difficult. Even the identifica

tion of the apparatus may offer problems, since the ele

ments partly cover each other. For example , when I studied

the type specimen of 'Paulinites ' polonensis Kielan:Jaworowska 1966, I had great problems in identifying the different elements composing the apparatus. It is a very small, probably juvenile, apparatus with indistinct morphological

details. The elements are fused together, slightly distorted

and fraetured. The study was further complicated since the

apparatus was kept in glycerine due to its fragile nature .

The decision to base the taxonomy on apparatuses more or

less forced Kielan:Jaworowska to choose a less good repre

sentative of the species as type, since apparatuses of paulin

itid taxa are rare .

Thus, the description of fused jaws in an apparatus is

complicated by the limited num ber of apparatuses and by

the fact that the elements partly hide each other. I believe

that it is an advantage to illustrate isolated elements as weU.

Reconstructions based on isolated elements have not

been very pop ul ar, yet there have been a num ber 'of more

or less successful attempts utilizing different methods. Syl-


Fig. 19. DA. Left MI and MIT fused together, Kel/nmtes (Kl aiJraham, Vattenfallsprofilen l, Hogk.lint b, 2 1 .4-2 1 .95 SL, ca. x l OO; Al lateral view, A2 lateral-dorsal view. DB. Left and right MI's and Mll ' s fused together, K sp. , slightly squeezed and fractured, LO 5761 : 1 , Vattenfallsprofilen l ; Hogklint Beds, unit b , 70-6L], x70; Bl lateral view, B 2 dorsal view. De. Dorsal view of right MI and left MIl fused together, Lanceolatites gracilis var. vis by, LO 5761 :3, Vattenfallsprofilen l , Hogklin t Beds, un i t b, 70-6L], xl OO. DO. VentraI view of left and right MI 's and MII 's fused together, K sp. , LO 5761 :2 , Vattenfallsprofilen l, HOgk.lint Beds, unit b, 70-6L], xl OO.

vester ( 1 959) attempted to reeonstruet an apparatus on the

basis of dispersed jaws. Although he did not support the

reeonstruetion by, e .g . , diversity or abundanee of different

elements, it might be eorreet. Later, Kielan:Jaworowska

( 1 966) reconstrueted some apparatuses from isolated elements, on the basis of eo-oeeurrenee of unusual element types ( left and right MI) .

Sueeessful reeonstruetions of apparatuses have also been made by Szaniawski ( 1 968) , on the Permian spe eies Atrakto

prion eudoxus, and by Corradini & Olivieri ( 1 974) , on the

Carboniferous-Permian species Brochosogenys siciliensis

(Corradini & Olivieri) Colbath 1 987b.

Reconstructing apparatuses from isolated jaws is a more

delicate task than analyzing more or less complete appara

tuses. However, dispersed jaws are easier to find and study,

and the greater number of specimens makes the choice of

a suitable type easier. As noted by Szaniawski & Gazdzicki

( 1 978) , reconstruction work of apparatuses is facili tated by

similarities between Recent forms and the fossil ones, espe

cially Mesozoic and Caenozoic . Ultrastructure also seems

useful for apparatus reconstruetion, and colour, surface

texture (e .g . pits and grooves) , and jaw size may be em

ployed, at least within the same sample .

Taxonomy based on quantitative studies of single ele

ments demands a large material, collected from localities

of different geographical areas and stratigraphical levels. In

theory, it is possible to encircle the jaws of a particular

species by studying the co-occurrence of many elements in

different environments. In practice, one usually has to work

with samples from a limited range of environments and

with a limited number ofjaws. Reconstructions may also be complicated by the presenee of severaI biological taxa in a

sample. Nevertheless, in most cases it has been possible to

reconstruct the apparatus by delimiting the taxa in 'favourable ' samples. In comparison with the limited number of apparatuses, the large number of isolated elements

makes it possible to choose types representing a mean

within the morphological variation.

In my opinion, synonymization of names between the

two taxonomical systems is inevitable, but must be done

very carefully. Literature studies alone are insufficient for

this purpose. A sound revision must be based on first-hand

studies of fossil material to determine apparatus composi

tion and variability of jaw morphology. The num ber and

form of the jaws forming an apparatus can be derived from

complete apparatuses or, at least in some taxa, numerical

studies (e .g . Corradini & Olivieri 1 974) . No less important

is the knowledge of the jaw variability within the popula

tion. Both ontogenetic and phylogenetic aspeets are signif

icant. A large number of specimens are required, and eaeh

element must be exposed so that a thorough study of every

part of the element can be carried out.


Fig. 20. Dorsal view of a reconstructed polychaete jaw apparatus of Kettnerites (Aeolus) sisyphi klasaardensis. The reconstruction with the elements separated includes the left and right of the following elements (from top) : MIV, left MIll, MIl, MI, and carriers. The reconstruction of the fused elements shows the position and the corresponding size of the left and right MI 's and MII 's in an apparatus.


Classification

Phylum Annelida Lamarek 1 809 Class Polychaeta Grube 1 850 Order Eunicida Dales 1963 Superfamily Eunicea Grube 1 852 Family Paulinitidae Lange 1947

Lange ( 1 947) described one speeies (Paulinites paranaensis)

within the family Paulinitidae . Speeies belonging to this

family are reported from North and South America and

Europe. The stratigraphical range of the Paulinitidae is not

known in detail, Kielanjaworowska ( 1 966, p. 1 25) gives the

following range : Ordovician to Carboniferous ( ?Permian) .

The paulinitid species from Gotland belong to the gen

era Gotlandites, Hindenites, Kettnerites, Lanceolatites and

Langeites. The criteria for delimiting the genera are in

short: The MI's of Kettnerites are fairly coarse with inner and

outer margins almost paralleI compared with Lanceolatites.

The MI and MIl of the latter genus are very slender, of ten

with a large number of denticIes, and the basal plate is

normally fused to the MI. Elements (MI's) belonging to the genus Langeites, so far monospecific , have only very few denticIes. The MI 's are slender with almost paralleI sigmoi

dal inner and outer margins ending anteriorly in a large

denticulated hook. The MI 's of Gotlandites and Hindenites

have a wide posterior part and a more pronounced distine

tion between the hook and the rest of the jaw. Gotlandites

differs from Hindenites in that the former has a larger hook

and a more pronounced convex inner margin. The MIl

elements also differ between these genera.

A key to the speeies is given in Table l .

Genus Gotlandites Bergman 1987

Derivation of name. - Latin Gotlandites, from the Island of

Gotland.

Type and only species. - Gotlandites slitensis Bergman 1987

40 Claes F. Bergman

Diagnosis. - Right MI: Jaw approximately triangular, taper

ing strongly towards the large fang, inner margin slightly

convex with paucidentate denticulation anteriorly. Denti

des decrease in size and become less spaced posteriorly;

large, wide basal portion, with a fairly small, fused basal

plate .

Left MI: Jaw approximately triangular, tapering anteri

orly, inner margin almost s-shaped, ending in a very large

fang with paucidentate denticulation on anterior half of

dentary.

Right MIl: Fairly large cusp with a small to a fairly small

pre-cuspidal dentide, followed by one or two minor denti

des. Large shank and large dentides.

Left MIl : Fairly large cusp without pre-cuspidal dentides,

four large intermediate dentides. Large, wide ramus.

Gotlandites slitensis Bergman 1987 Figs. 140, P, 1 8N, 21 , 22

Synonymy. - O 1987 Gotlandiles slilensis n.sp. - Bergman, pp.

34-36, Figs. 1 40, P, 18 , 2 1 , 22 .

Derivation of name. - Latin slitensis, from the Slite Beds.

Holotype. - LO 5780 :5, left MI, Fig. 21 G.

Type locality. - Vike 2.

Type stratum. - Slite Beds, Slite Mari , Penlamerus gothlandicus

Beds or slightly older.

Material. - Fig. 3; About 44 right MI, 42 left MI, 64 right MIl,

and 41 left MIl .

Occurrence. - Figs. 4 and 6; Wenlock. Slite Beds, unit g and

Slite MarI . Ajstudden l , Fardume l , Hide l, Hide Fiskelage

l , Munkebos l , Nygårds l , Slitebrottet l , Talings l , Tjel

dersholm l , Vallstena 2, Vike l , 2, and 3.

Diagnosis. - As for the genus.

Description. - Right MI, dorsal side: Length 0.38-1 .85 mm; width about 0.3 times the length . The jaw is approximately triangular and tapers towards the strongly developed fang. The inner margin is convex, denticulated from the falcal arch to the undenticulated ridge . The slightly slanting dentides occupy about 0 .5-0.6 of the total jaw length in smaller specimens, and about 0 .6-0 .7 in larger ones ( i .e . arou.nd 1 .5 mm or longer) . The anteriormost 4-5 dentides are moderately large to large , and paucidentate to widely spaced. The posterior remaining 5-1 0 dentides are dosely spaced, ending abruptly at the undenticulated ridge . In smaller jaws the denticulation en ds before reaching the inner wing; in larger ones it continues posteriorly, ending beside the basal plate . The length of the undenticulated ridge represents about 0 . 1 5-0.30 of the total length in smaller jaws and about 0 . 1 0-0 . 1 5 in larger ones. The undenticulated ridge forms the posterior sharp tip of the jaw, though in some cases the ridge ends without reaching the posteriormost part. The inner wing is almost rhomboidal, normal-sized; the inner margin parallei to the undenticulated ridge . The normal basal furrow is partly hidden under the skew, rectangular, fused basal plate . The poste-


rior margin is almost straight, directed slightly anteriorly,

and forming a smoothly rounded outer margin together

with the outer margin of the large basal portion. The basal

angle is about 30-35° . A more or less pronounced rim runs

along the anterior outer margin of the basal portion . The

outer margin anterior to the basal portion is smooth,

shaped almost like a reversed s.

Ventrai side : The crescent-shaped myocoele opening is

narrow, strongly endosed, representing about one third of

the jaw length . The opening is surrounded by a narrow rim,

and part of its interior surface forms a wide, smooth,

rounded area along the outer margin .

Left Ml, dorsal side: Length 0.35-1 .93 mm, width 1f4-1f3

of the length . The subtriangular, strongly taperingjaw ends

in a slender, very large fang. The inn er margin is s-shaped.

The number of dentides is coupled to jaw size, the smaller

jaws having 4-5 smaller, pronouncedly paucidentate denti

des, and the larger ones up to around 20 dentides. The

dentides are of normal size (cf. Fig. 15 ) and become sub

dued posteriorly, in larger jaws ending as crenulations. In

larger jaws the size and spacing of dentides varies, with one

or a few dosely spaced, very small dentides interrupting a

series of dentides of decreasing size and space. The ante

rior dentides are indined slightly towards the anterior. The

denticulation occupies around 0.3 to more than 0 .6 of the

length of the inner margin in smaller and larger jaws,

respectively.

The transition from juvenile to adult is gradual . When

the jaw attains a length of about 1 .5 mm the num ber of

dentides starts to increase , and the shape begins to change

from sickle-shaped to more triangular.

The transition from the rounded denticulated inner

margin to the undenticulated ridge is smooth . The inner

wing is of normal size, about 1f4 of the jaw length, and the

rounded inner margin follows the orientation of the un

denticulated ridge . The basal furrow is small, with a large

outer face . A more or less pronounced furrow anterior of

the outer face, dividing it from the rest of the jaw, is not

uncommon; the structure resembles a very fused basal plate . The basal angle is much varied. The anterior part of the outer margin to the basal portion forms an outer wing.

The outer margin anterior of the basal portion is smoothly

rounded, of ten with a concavity in the lower middle part,

ending in a large, sickle-shaped falx.

Ventrai side : Almost a mirror image of the ventrai side of the right MI.

Fig. 21. Gotlandiles slilensis. All speeimens except l and K from Slite Beds, Slite Mari, Penlamerus gothlandicus Beds or slightly younger. Speeimens A, B, D, E, G, H,] , L and M are from Vike 2, 83-4GB. All speeimens are in dorsal view. DA. Left MIl, LO 5780 : 1 , x 1 20 . DB. Right MIl , LO 5780:2 , x1 20. D G. Right MlV, LO 5779: 1 , Ajstudden l, 83-7GB, x1 70. DD. Left MIl, LO 5780:3, x80. D E. Right MIl, LO 5780:4, x80. DF . Vike l, 75-30GB; Fl left Ml, LO 5785 : 1 , x60; F2 right MI, LO 5785:2 , x60; F3 left MIl, LO 5785:3, x60; F4 right MIl, LO 5785 :4, x60. DG. Holotype, left MI, LO 5780:5, x80. DH. Right MI , LO 5780:6, x1 20. DI . Right MI , LO 5774:2 , Slitebrotte t l , Slite Beds, Slite Mari, 73-37L], x 1 20. Dj. Right MI, LO 5780:7, x80. DK Left MI, LO 5774 : 1 , same sample as I, x150. D L. Left MI, LO 5780:8, x1 20. DM. Right MI and MIl fused together, LO 5781 :3, x 1 70.

FOSSILS AND STRATA 25 ( 1 989) Silunan paulinitid polychaetes from Gotland 41


Fig. 22. Gotlandites stitensis. Dorsal (A-F) and ventrai (G-J) views. DA. Left MI, LO 5776: 1 , Vallstena 2 , Slite Beds, Slite Marl Pentamerus gothlandicus Beds or slightly younger, 77-2CB, x80. D B. Left MIl, LO 5779:2 , Ajstudden l , Slite Beds, Slite Marl Pentamerus gothlandicus Beds (?) , 83-7CB, x 170. De. Right MI, LO 5779:3, same sample as B, x80. OD. Right MIl, LO 5839:4, Gandarve l , Halla Beds, 7 1 -84LJ, x 1 20. DE. Left

MI, LO 5778:3 , Munkebos l , Slite Beds, Slite Marl, Pentamerus gothlandicus Beds or slightly younger, 7 l-84LJ, x120 . O F. Right MI, LO 5776:2, same sample as A, x 1 20. D G-J. Vike 2 , Slite Beds, Slite Marl, Pentamerus gothlandicus Beds or slightly younger, 83-4CB, ventrai views, x1 20; G, left MI, LO 5805:3 ; H, right Mil, LO 5805:4; l, left MIl, LO 5805:5 ; J, left MIl, LO 5805 :6.


Right MIl , dorsal side : Length 0.32-1 .50 mm, width

about half the length . A small to moderately small pre

cuspidal denticle forms the fairly sharp anteriormost part

of the jaw, somewhat variably situated on the cusp . The

cusp is of normal size, slightly swollen basally, slightly

slanted, and followed by one or two minor denticles form

ing the intermediate dentary. The post-cuspidal dentary is

large, composed of 6-8 slanted denticles which decrease

posteriorly in size. The shank is large, about 0 .6 of the jaw

length, posteriorly tapering, with inner margin alm ost

straight to slightly convex, outer margin slightly concave,

and posterior margin rounded. The bight angle is difficult

to measure due to the variable ramus, which is fairly short,

sigmoidal, wide, and more or less blunt-ended with a

smaller sinus in the anterior outer margin.

Ventrai side : The slightly enclosed myocoele opening

represents about �3 of the jaw length . A fairly wide rim

forms the anterior margin of the myocoele opening and

continues along the ramus; it also extends along the shank.

The shape of the margins varies considerably.

Left MIl, dorsal side : Length 0.43-1 .43 mm, width about

half or slightly more than half of the length . The cusp is of

normal size, and slightly swollen in its basal part. The

intermediate dentary is composed of four denticles: three

large denticles of equal size and a posterior, slightly larger

one. There are 6-8 large , slanting, post-cuspidal denticles,

decreasing in size towards the posterior. The shank occu

pies about half the length of the jaw, tapers posteriorly, and

has a rounded ending. The almost triangular inner wing is

widest an teriorly and occupies about half the length of the

jaw. The bight angle is acute , the ramus is large, wide, and

tapering, with a fairly rounded end having a distinet sinus

at the anterior outer margin.

Ventrai side: The slightly enclosed myocoele opening

extends from slightly more than half to two thirds of the jaw

length . The moderately ereseent-shaped anterior margin

of the myoeoele opening extends along the ramus.

Comparisons. - The morphology of the MI is unusual ,

though the corresponding elements of the Late Palaeozoic

Brockosogenys siciliensis (Corradini & Olivieri 1974) Colbath

1 987b show a faint general resemblance in having a large

fang and similar dentieulation . The MII ' s are different in the two speeies. Due to the large time span separating B. siciliensis and G. slitensis, their relationship must remain

hypothetieal.

The left MIl of Gotlandites slitensis eould be mistaken for

a left MIl of Hindenites gladiatus. G. slitensis has a larger

ram us and usually a smaller pre-eusp than H. gladiatus.

Genus Hindenites Bergman 1 987

Derivation of name. - In honour of G. J. Hinde , one of the

pioneers of the study of fossil polyehaetes and the first to

publish a paper on the polyehaete jaws from Gotland.

Type species. - Hindenites angustus (Hinde ) .

Otker species. - H. gladiatus, H. naerensis.

Silurian paulinitid polyckaetes from Gotland 43

Diagnosis. - Right MI: Jaw tapers anteriorly, ending in a

strong fang. Convex inner margin, fairly large denticles.

Myoeoele opening on ventrai side extending for about

0.35-0.5 of length ofjaw.

Left MI: More or less a mirror image of righ t MI exeept

for basal portion. Ioner wing almost triangular.

Right MIl: Slender, very narrow ramus, long shank with

almost parallei inner and outer margins. Cusp indistinet.

Left MIl: As for right MIl, exeept for cusp whieh is more

pronouneed but still fairly small.

Hindenites angustus (Hinde 1 882) Figs. 14I , 1 80, 24B: 1-7

Synonymy. - O 1 882 Arabellites angustus n. - Hinde, p. 19, Pl.

2:53, left MIL 0 1 882 Arabellites arcuatus n. - Hinde, p. 19 ,

Pl . 2 :52, right MIl . 0 1 980 Paulinites gladiatus Kielan-:Jawor

owska 1 966 - Wolf, pp. 86-87, Pl . 1 2 : 1 05-1 07. 0 1 987 Hin

denites angustus (Hinde 1 882) - Bergman, pp. 37-40, Figs.

1 41, 18 , 24B: 1-7.

Holotype. - Left MIl, British Museum A 222 1 , illustrated in

Hinde 1 882, p. 19 , Pl. 2 :53.

Type locality. - Vattenfallsprofilen l , Gotland.

Type stratum. - Hogklint Beds, unit d, 'Pterygotus ' mari,

Herrmannina Beds.

Material. - Figs. 2 and 3; at least 60 right MI, 59 left MI, 39

right MIl, and 40 left MIl.

Occurrence. - Figs. 4 and 1 1 ; Early to Middle Wenloek.

Hogklint, Slite, and Klinteberg Beds. Ansarve l , Follingbo

12 , Klinteberget l , Loggarve 2 , Strandakersviken l , Vatten

fallsprofilen 1 .

Diagnosis. - Right MI: Tapering jaw with strong fang, eonvex inner margin, dentieulated from middle part of falcal

areh down to undentieulated ridge . Shank triangular and

pointed, outer margin almost straight.

Left MI: Taperingjaw with eonvex inner margin, dentie

ulated from extreme anterior end of falcal areh ending

before reaching the short and almost rectangular inner

wing. Basal furrow short and oriented parallei to outer

margin of basal portion. Right MIl: Small double eusp, shank long and slender,

ramus long, narrow and pointed. Left MIl: Cusp followed by three intermediate denticles

and one large eusp-like denticle. Shank fairly long, ramus pointed.

Description. - Right MI, dorsal side : The jaw is 0 .29-1 .08 mm

long, variable in width (usually about V3 of the length) ,

tapering strongly from lower middle part. Anteriorly it

ends in a large fang, usually bent upwards with regard to

the fairly rounded dorsal surfaee . The inner margin is

convex and 0 .6-0.7 the length of the jaw. Dentieulation

extends from the anterior part of the faleal areh to a

position anterior to the inner wing. The slightly slanting

denticles increase somewhat in size towards their maxi

mum in the posterior middle part; in some jaws the largest

denticles are anterior. The posteriormost denticles usually

44 Claes F. Bergman

gradually decrease in size and end abruptly. The num ber

of denticles varies from 1 2 , for ajaw length of 0 .40 mm, to

19 , recorded on 0.53-1 .08 mm long jaws. The undentic

ulated ridge is long, about 0.3 of the jaw length, straight

and fairly narrow; the transition to the denticulated ridge

is smooth. The inner wing is nearly triangular with the

anterior inner ' corner' sickle-shaped, of normal size (0 .25

of the jaw length) , strongly downfolded, almost perpendic

ular in relation to dorsal surface . The posteriormost part of

the inner wing forms, together with the undenticulated

ridge, the moderately sharpened posterior part of the

shank. The basal furrow is long, fairly narrow, extending

from the posteriormost denticles down to the posterior end

of the shank.

Basal plate not recorded.

The flange is moderately thick-walled, highly elevated in

the posterior, strongly downfolded along the long practi

cally straight, outer margin. The ligament rim is exposed as

a narrow straight ridge along the outer anterior margin of

the basal portion. The bight angle is wide , about 1 00° , and

the basal angle very sharp, about 5 0. Thus, the ligament rim

and the undenticulated ridge are almost paralleI. The

outer margin anterior to the basal portion is almost straight

with only a minor median concavity, and ends in a large

sickle-shaped falx.

Ventrai side : The enclosed crescent-shaped myocoele

opening extends anteriorly for about 0.4-0 .5 of the jaw

length . The opening is surrounded by a thickened rim

along the anterior and outer margins.

Left MI, dorsal side : Length 0.39-1 . 1 4 mm, width 1J4-1J3

of the length . The jaw tapers anteriorly, ending in a large

fang, which is bent upwards in relation to the dorsal sur

face. The inner margin is convex, and the denticulation

extends from the anterior part of the falcal arch over about

0.6 of the jaw length, ending short of the inner wing.

Denticle num ber varies more or less regardless of size from,

14 denticles on a 0 .48 mm longjaw to 24 on one of 0 .64 mm

length, but 18 and 20 denticles have been recorded onjaws

0 .39 mm and 1 . 1 4 mm long, respectively. The anteriormost denticle is the largest, normally followed by some denticles decreasing in size towards the posterior part of the falcal

arch, from where the denticle size increases to fairly large

ones on the middle part of the inner margin. The poster

iormost denticles are small. In some jaws the denticulation gradually decreases in size from the large st, anteriormost

denticle. The anterior denticles point forward while most

of the remaining denticles are slanting. The inner margin

continues as a straight line onto the small and posteriorly

narrowing undenticulated ridge. The moderately down

fold ed inner wing is angular, nearly rectangular, with an

almost straight inner margin, representing about 0 .2 of the

jaw length . The basal furrow is short and narrow, parallei

with the almost straight outer margin of the basal portion .

The outer face of the basal portion is large , slightly more

than 0.3 of the jaw length . The basal angle is narrow, about

15 ° . The outer margin anterior to the basal portion is

slightly convex with a smaller concavity in the midd le part.

Ventrai side : The enclosed myocoele opening is crescent

shaped anteriorly, directed toward the anterior, sur-


rounded by the fairly low ligament rim, which extends

posteriorly at the inner and outer margins in the form of

narrow spur-like ridges.

Right MIl, dorsal side : Length 0 .45-0 .80 mm, width

about �3 of the length. There is a very small double cusp,

anterior cusp slightly smaller than the posterior one. They

are followed by one, intermediate , fairly small denticle , in

turn followed by seven to eight post-cuspidal denticles, only

slightly smaller than the cusps. The denticles are slanting

and decrease in size towards the posterior, ending before

they reach the the posterior margin . The inner margin is

slightly convex. The shank is extremely long, occupying

more than two thirds of the length of the jaw, and the fairly

slender and posteriorly pointed tip is somewhat bent to the

outside . The margins of the shank are almost parallei (Fig.

24B:5) or taper slightly posteriorly (Fig. 24B:7) . The small

inner wing is normally not seen from the dorsal side . The

bight is deep, the bight angle about right to acute . The

ramus is very slender, with almost parallei margins and a

pointed extremity. The outer margin is of ten slightly con

vex. A minor sinus is sometimes present on the anterior

outer margin .


represents about 3f4 of the jaw length . It is surrounded by a

very narrow rim which forms a protruding corner on the

anterior part of the inner margin.

Left MIl, dorsal side : Length 0.38-0.85 mm, width about

half the length . One cusp of normal size forms the sickle

shaped anterior margin. It is followed by three fairly small

intermediate denticles which increase posteriorly in size. A

conspicuous, large, cusp-like denticle divides the interme

diate from the post-cuspidal dentary. There are eight to

nine post-cuspidal denticles, about equal in size and slightly

slanting towards the posterior. The almost parallel-sided

shank is very large , about 0 .6 of the jaw length, ending fairly

bluntly. The posteriormost part extends to the outer ven

tral side . The bight is very deep, and the bight angle sharply

acute . The ramus is fairly long and slender with a posteri

orly oriented, pointed extremity. The anterior outer margin is almost straigh t.

Ventrai side : The slightly enclosed myocoele opening represents about two thirds of the jaw length. The rim

which surrounds the opening to the anterior ( slightly

curved) and along the inner margin of the jaw forms an alm ost right angle.

Discussion. - The holotype is the left MIl; the right MIl of

the same species was described by Hinde ( 1 882) from

Gotland as 'A '. arcuatus. A North American right MIl with a

similar denticle formula was described by Hinde ( 1 879) as

'A '. similis var. arcuatus. Although I have not seen the North

American specimen, there seems to be a distinct difference

between the Gotland form and the jaw from the North

American Middle Devonian Hamilton Group. 'A '. similis

(Hin de 1 879) from the Silurian Niagara Formation in

North America also has a similar denticle formula, but

judging from the crude illustration and description there is

no cio se relationship to the Gotland form. Hinde did not

com pare this form with the one from Gotland ( 'A '. arc

uatus) .


Companson. - The large myocoele opening of the MI is

characteristic of the genus, distinguishing it from other

paulinitid genera on Gotland. The MI differs from the

corresponding element of Hindenites gladiatus in that the

denticulation en ds before reaching the inner wing. The

MIl has a unique dentition and form.

Hindenites gladiatus (Kielan:Jaworowska 1966) Figs. 1 4F, G, H, 1 8P, 23, 24A, C-E

Synonymy. - 0 1 882 Arabellites kamatus Hinde 1 879 - Hinde,

pp. 1 6-1 7 (pars) , Pl. 2 :42-43, left MI. 0 1 966 Paulinites

gladiatus n.sp . - Kielan:Jaworowska, pp. 1 29-1 3 1 , Pl. 30:5A

C, 6A-C, left and right MI. 0 1979 Oenonites sp. a - Berg

man, p. 99, Pl. 28 :3A, B, left and right MI. 0 1 987 Hindenites

gladiatus (Kielan:Jaworowska 1 966) - Bergman, pp. 40-42,

Figs. 1 4F-H, 18, 23, 24A, C-E.

Holotype. - Right MI in Kielan:Jawbrowska 1 966, pp. 1 29-

1 3 1 , Pl. 30:6.

Type locality. - Loose boulder derived from the Baltic area.

Type stratum. - Silurian (?Upper Wenlock, ?Lower Ludlow) .

Material. - Figs. 2 and 3; At least: 83 right MI, 55 left MI, 26


Occurrence. - Figs. 4 and I l ; Early Wenlock to Middle-Late

Wenlock, H6gklint Beds, unit b, to Mulde Beds. Ar l ,

Aursviken l , Dapps 2 , Fardume l , Gandarve l , Hallagrund

l , Hide l , Munkebos l , Nygårds 2, Saxriv l , Slitebrottet l

and 2, Svarven l , Valby Bodar l , Vallstena 2, Vattenfallspro

filen l , Vike 2 .

Diagnosis (emended) . - Right MI: Large denticles covering

inner margin usually from posterior of the falcal arch, passing posteriorly to anterior part of the inner wing, end-

Fig. 23. Hindenites gladiatus (Kielan-jaworowska 1 966) , eamera-lueida drawing of the type specimen (right MI, Z. Pal. No. O.308/ 1 4b) and the left MI of the same speeies, x84. Note that the denticulation of both right and left MI ends slightly to posterior of the anteriormost part of the inner wing.

Silunan paulinitid polyckaetes from Gotland 45

ing slightly anterior of the bight. Myocoele opening cover

ing 0.35-0.40 of the jaw length .

Left MI: Large fang, falcal arch with pronounced cutting

edge but without true denticles . Large denticles along

convex inn er margin, end posterior of the anterior part of

the inner wing.

Right MIl: Fairly small pre-cuspidal denticle, cusp of

normal size, very large shank, ram us long, narrow, with

pointed extremity.

Left MIl: Fairly large single cusp, very large shank, long,

very narrow ramus with pointed extremity.

Description. - Right MI, dorsal side : Length 0.55-l .20 mm,

width V4-V3 of the !ength . The jaw tapers strongly anteri

orly, ending in a large, pointed, sickle-shaped fang, bent

upward in relation to the jaw surface . The denticulation of

the an terior part of the inner margin is forrned by more or

less indistinctly spaced crenulations, or as in the specimens

from Saxriv l (H6gklint Beds, unit b) , by a cover of small

denticles. The anterior denticles seem to be more devel

oped in the smaller jaws, at !east during Middle-Late Wen

lock time. In the Slite MarI at Munkebos l an excellently

preserved jaw exhibits large, thin, triangular denticles an

teriQrly. In the convex posterior part of the inner margin,

the 1 1-13 denticles are large and slanting, passing along

the anterior part of the inner wing, close to the bight. In

the larger jaws from the Early Wenlock the posteriormost

3-5 denticles decrease in size towards the undenticulated

ridge . In Middle-Late Wenlock specimens only the last

denticle is smaller (e.g. Slitebrottet, Munkebos l , and

Gandarve l ) . Denticles of the smaller jaws are of more or

less equal size on the posterior half of the denticulated

inner margin. Early Wenlock jaws show a prominent, short

and straight undenticulated ridge . In Middle-Late Wen

lock specimens the ridge is somewhat broader, the broad

est part being at the posterior end. The strongly down

folded, almost triangular, inner wing occupies about 0 .3 of

the jaw length . The basal furrow has a smooth outer side

and is wide and long, about twice the length of the un

denticulated ridge .

Basal plate is not recorded.

The flange is thick-walled, bight angle about 90° . The

basal angle is low, varying between 1 2° and 22° . The ligament rim is almost straight, occupying around one fifth to one quarter of the jaw length . The outer margin in front of the basal portion is convex with a more or less pronounced

median concavity. The falx is large , wide, sickle-shaped.

VentraI side : The enclosed myocoele opening occupies

around 0.35-0.40 of the jaw !ength . The opening is cres

cent-shaped anteriorly and surrounded by a sharp ridge

along the innermost part of the inner wing.

Left MI, dorsal side : Length 0.53-l .26 mm, width about

V4 of length. The jaw tapers strongly anteriorly, ending in

a large , pointed, sickle-shaped fang. The fang has a promi

nent undenticulated cutting-edge along the large falcal

arch, com prising 0 .3-0.45 of the anterior part of the jaw,

though 2-8 indistinct knobs may occur. The posterior part

of the inner margin is convex, with 9-1 5 large to very large

slanting denticles, occupying 0 .34-0.50 of the jaw length .

The largest denticles are in the middle part of the convex


Fig. 24. All speeimens in dorsal view. DA. Hindenites gladiatus, Vattenfallsprofilen l, Hogklint Beds, unit d, Valdana testudo bed, RM sample ; Al left MI, AN 2702, xSO ; A2 right MI, AN 2703, xSO; A3 left MIl, AN 2704, xSO; A4 right MIl, AN 2705, xSO; A5 left Mn, AN 2700, xSO; A6 right Mn, AN 270 1 , xSO. DB. H. angustus, Vattenfallsprofilen l , Hogklint Beds, unit d, Herrmannina bed, RM sample ; Bl left MI, AN 2706, x60; B2 left MI, AN 2707, xSO ; B3 right MI, AN 270S, xSO; B4 left MIl, AN 2709, x80; B5 right MIl, AN 2710, xSO; B6 left MIl, AN 27 1 1 , x1 20;

B7 right Mn, AN 2712, xSO. DC. H. gladiatus, Munkebos l , Slite Beds, Slite Mari, Pentamerus gothlandicus Beds or slightly younger, 71 -84Lj; Cl left MI, LO 577S : I , x1 20; C2 right Ml, LO 5778:2 , x1 20. OD. H. gladiatus, Gandarve l , Halla Beds, 7 l-8ILj; D l left Ml, LO 5S36:6, x1 20; D2 right MI, LO 5S36:7, x 120 . DE. H. gladiatus, right MI, LO 5774: 3, Slitebrottet l , Slite Beds, Slite Mari, 73-37Lj, x1 20.


inner margin. The undenticulated ridge is short, tapering

anteriorly, forming a more or less straight continuation of

the inner margin. At the posterior end it ben ds toward the

base of the inner wing, which occupies about lf4-lf3 of the

jaw length . Inner wing varies from nearly triangular to

semicircular and, not so deeply downfolded as in the right

MI, forms the posteriormost part of the jaw. The basal

furrow is almost triangular and short, tapering anteriorly,

its length about half the length of the inner margin. The

almost straight ligament rim is notable from the dorsal view

and occupies about one third of the jaw length. The basal

angle is low. The outer margin in front of the basal portion

is smoothly convex with a pronounced cutting-edge in the

falx, though sometimes a shallow concavity in the middle

of the jaw may occur.

VentraI side: The endosed myocoele opening extends

for 0.35-0.40 of the jaw length . The opening is crescent

shaped anteriorly and the ligament rim is thickened and

surrounded by a narrow furrow on the anterior and inner

part.

Right MIl, dorsal side : Length 0.45-0.78 mm, width

about % of the jaw length . A small pre-cuspidal dentide,

on the cusp, forms the anteriormost, fairly rounded mar

gin . The inner margin is slightly convex. The cusp is of

normal size, followed by the post-cuspidal dentary. The

anteriormost dentide is slightly smaller than the following,

about 8, large dentides, which slant somewhat towards the

posterior. The shank with almost parallei sides is very large,

and occupies about 0.6 of the jaw length; the posterior

extremity is fairly narrow and slightly rounded. The bight

is deep, the bight angle acute to almost 90° . The ram us is

long, very narrow, with a posteriorly oriented, pointed

extremity. The anterior outer margin has a slight sinus.

VentraI side : The slightly endosed myocoele opening

represents about % of the jaw length. It is delimited by the

smooth rim along the almost straight anterior and inner

margins.

Left MIl, dorsal side: Length 0.36--0.88 mm, width ap

proximately �3 of the jaw length . The inner margin is

slightly convex. A single cusp of normal size is followed by

the intermediate dentary containing 3-5 dentides of nor

mal size, increasing slightly towards the posterior. The

tra·nsition to the post-cuspidal dentary is more or less grad

ua! . The size of the post-cuspidal dentary is varied, from normal to fairly large , posteriorly slanting dentides. The

denticulation continues along the entire shank, which is

very large, paraIlel-sided, and occupies about 0 .6 of the jaw

length; the posterior extremity ends bluntly. The inner

wing is very short and has a rounded margin . The bight is

fairly deep, the bight angle acute . The parallel-sided ram us

is conspicuously long and slender, with a pointed extremity

oriented towards the posterior. A pronounced sinus is

found on the anterior outer margin.

Ven tral side : Almost a mirror image of the ven tral side of

the right MIl, except that the opening is more rounded

and the inner wing extends to the rim on the left MIl.

Remarks. - The most characteristic difference between H. gladiatus and H. angustus is that the dentides of the right

MI of H. gladiatus extends, on the denticulated ridge, along


the anteriormost part of the inner wing. The conspicuously

large dentide of the left MIl of H. angustus is also a distinct

character.

Comparisons. - The type species Arabellites kamatus Hinde

( 1 879, p. 377, Pl. 1 8 : 1 2) was found in the North American

Cincinnati Group. The type specimen is squeezed and not

well preserved, but it does not belong to the same species

as the specimens described as A. kamatus from Gotland

(Hinde 1 882, pp. 1 6--1 7, Pl. 2 :42-44) .

Kielan:Jaworowska ( 1 966, pp. 1 30-1 3 1 ) noted that Nerei

davus digitus Eller ( 1 963, p. 1 6 1 , Pl. 1 : 1 4) from the Upper

Devonian Sheffield Shale of Iowa and Nereidavus exploratus

Eller ( 1 96 1 , p. 1 77, Pl. 1 :20-2 1 ) from the Devonian of

Michigan were 'reminding one in the shape of Paulinites

gladiatus n.sp. ' . In my opinion it is difficult to discern

whether or not there is a doser relationship between the

Baltic and the Laurentian taxa without studying popula

tions from North America.

Hindenites naerensis Bergman 1987 Figs. 1 4] , 1 80, 25

Synonymy. - O 1 987 Hindenites naerensis n.sp. - Bergman, pp.

42-44, Figs. 1 4J, 18 , 25.

Derivation of name. - From the parish of Nar, referring to the

type locality.

Holotype. - LO 5831 :4, right MI, Fig. 25B.

Type locality. - Narshamn 2.

Type stratum. - Burgsvik Beds, lower part.

Material. - Fig. 3 ; more than 24 right MI, 26 left MI, right

MIl 5, left MIl 6.

Occurrence. - Figs. 4 and 1 1 ; Early Whitdiffian, Ludlow.

Burgsvik Beds. Glasskar l , Kapelludden l , Narshamn 2 .

Diagnosis. - Right MI : Jaw smoothly rounded, tapering

anteriorly, conspicuous almost triangular inner wing.

Large , crescent-shaped, endosed myocoele opening.

Left MI: Jaw smoothly rounded, tapering anteriorly,

large crescent-shaped endosed myocoele opening.

Right MIl: No distinguishable cusp or intermediate dentary, shank extremely long, occupying about three quarters

of the jaw Iength . Ramus very narrow and slender.

Left MIl: Fairly small cusp, small post-cuspidal dentides.

Shank narrow with parallei sides and extremely long, occu

pying about three quarters of the jaw length .


width lf4-lf3 of the length . The jaw tapers unevenly anteri

orly, and the fang is slightly bent upward in relation to jaw

surface. The inner margin is somewhat convex, with cren

ulations of up to 5 small dentides along the inner margin

of the faJcal arch. Behind the faJcal arch, there are 9-1 6

fairly large, slanting, and somewhat spaced dentides. The

denticulated inner margin covers about 0.6 of the jaw

length. The posteriormost dentides decrease in size poste

riorly. The undenticulated ridge is long, 0 .3 of the jaw

length, narrow and low, forming a smooth arch. The ridge

48 Claes F. Bergman

Fig. 25. Hindenites naerensis. Specimens A, B and D in dorsal view. Narshamn 2, Burgsvik Beds, 83-1 2LJ; DA. Left MI, LO 5831 :3, x80; D B. Holotype, right MI, LO 5831 :4, x80. D e. Right MI, ventraI view, LO 5805 :7, x 1 20. OD. Right MIl, LO 5805:8, x80.

disappears gradually without reaching the posterior margin . The inner wing is anteriorly rounded, short, about V4

of the jaw length, strongly downfolded in comparison with

the dorsal surface, and forms the blunt posterior termina

tion of the jaw. The basal plate is small, nearly rectangular, without den

ticles. The plate and adjacent area of the outer face are highly elevated, forming the highest part of the jaw, the

fang excluded, making the posterior margin undulating.

The outer margin of the basal portion is downfolded and

almost straight in anterior-posterior direction. The basal

furrow is wide, the deepest part being between the anterior

most part of the basal plate and the undenticulated ridge .

The very long iigament rim extends almost 0 .4 of the jaw

length , along the outer margin. The basal angle is very low,

the undenticulated ridge and the outer margin are almost

paralleI. The outer margin is almost straight, the basal

portion included, with a minor convexity in the middle part

of the jaw.

VentraI side : The elongated, enclosed myocoele opening

represents about half of the jaw length . The opening is

surrounded by a smooth, somewhat sunk ligament rim. A


ridge , corresponding to the boundary between the basal

plate and the flange on the dorsal side of the jaw, runs

perpendicularly to the long axis of the jaw.

Left MI, dorsal side : Length 0.62-1 .53 mm, width about

lf4-lf3 of the jaw !ength . The jaw tapers anteriorly, ending

in a fang, bent slightly upwards in relation to the dorsal

surface. The inner margin behind the undenticulated

falcal arch is convex with some minute , and about 1 4-17

larger, slightly spaced, slanting denticles covering about

half the length of the jaw. The denticle size increases from

the anterior to the middle portion where it reaches a

maximum and then decreases posteriorly. The undentic

ulated ridge is long, low and narrow with a small median

inward bend, and normally does not reach the posterior

margin. The inner wing is of varying but always small size,

nearly rectangular and moderately downfolded in relation

to the dorsal surface. The basal furrow is variable , but

usually very short, almost rectangular, with its widest part

posteriorly. The outer margin of the basal portion extends

0 .3-0.4 of the jaw length, is slightly convex, deeply down

folded, almost hiding the long ligament rim. The outer

margin in front of the basal portion is mostly occupied by

the large sickle-shaped falx.

VentraI side : The enclosed, elongated myocoele opening

extends for approximately 0.4 of the jaw length. The liga

ment rim is fairly thin, except at the outer anterior position

where it forms a broad thick rim, narrowing posteriorly.

The posteriormost part of the rim forms a narrow, sharp

ridge .

Right MIl, dorsal side : Length 0 .77-1 .00 mm, width

slightly less than �3 of the jaw length . No cusp or interme

diate denticles are distinguished. The number of denticles

varies between 9 and 13 without relation to jaw size. They

cover the entire length of the jaw and slant slightly towards

the posterior. The size decreases slightly to the posterior,

with the exception of the much smaller posteriormost den

tide. The very narrow and extremely long (about 3f4 of the

jaw length) shank has almost paralleI sides and a slightly

convex inner side . It occupies about three quarters of the

jaw length . The posterior extremity is slightly widened and bent somewhat ventrally outwards. The bight is very deep, the bight angle acute , the ramus long and very narrow with

a pointed extremity. The anterior outer margin is normally

slightly convex to almost straight.

VentraI side : The slightly enclosed myocoele opening

represents about 3/4 of the jaw length or more . It is boarded

by a fairly narrow rim, gently crescent-shaped along the

anterior and inner margins.

Left MIl, dorsal side : Length 0.71-1 . 1 8 mm, width about

half of the jaw length. A fairly small cusp forms the rounded

anterior margin. The intermediate dentary has 1-3 small

der'lticles increasing posteriorly in size. Ten to eleven fairly

small post-cuspidal denticles decrease evenly in size from

the base of the cusp to the posteriormost extension of the

jaw. The shank, with almost paralleI sides, is extremely long,

occupying about 3f4 of the jaw length . The posterior ex

tremity is slightly swollen, bent somewhat to the ventraI

outer side . The inner wing is of normal length, about 0 .6 of

the jaw length, and very narrow, the margin parallel with


the denticulated ridge . The bight is deep, the bight angle

acute, the ramus very slender with pointed extremity. The

anterior outer margin has a slight sinus.

Ventrai side : A mirror image of the ventrai side of the

right MIl except that the opening represents about �3 of

the jaw length .

Remarks. - The difference in the num ber of MI and MIl

encountered is probably due to the greater fragility of MIl,

but also to the fact that the MI is larger and therefore easier

to recover. MI is also more easily identified by fragments.

Comparisons. - MI differs from the corresponding element

of H. gladiatus by be ing more slender and straight, and in

that the right MI has a fused basal plate . The dentiele

formula of the MIl ' s is similar in the two species, but the

latter is slightly more slender, particularly the ramus, than

the former one . The denticulation of the MI ' s of H. naer

ensis and H. angustus are similar. H. naerensis is probably

more elosely related to H. gladiatus than to H. angustus.

This assumption is based on the fact that the dentition

formula of MIl seems to be less changed through time. H.

gladiatus and H. naerensis may even be in the same lineage .

Genus Kettnerites Zebera 1935

Jaw elements, found in Palaeozoic rocks from Europe and

herein referred to Kettnerites, have been placed in the fol

lowing genera: O Oenonites Hinde 1 879 ( O. infrequens Hinde

1 879, p. 382 Pl. 20:2 , left MI; O. compactus Hinde 1 879, p.

384, Pl. 20: 1 3, left MIl; O. aspersus Hinde 1 880, p. 373, Pl.

1 4: 7,8 , left and right MI; Hinde 1 882, p. 13 , Pl. 1 :2 1 , 22, 22A) . O Nereidavus Grinnell 1 877, (Nereidavus solitarius

Hinde 1 879, p. 385, Pl. 20: 12 , left MI) . O Kettnerites Zebera

1 935 (Snajdr 1 95 1 ; Mannil & Zaslavskaya 1985a and

1 985b) . O Arabellites Hinde 1 879 (A. perneri Zebera 1 935) .

O Pronereites Zebera 1935. OPaulinites Lange 1 947; the type

species from the Devonian of South Ameriea belongs to a

different genus from Kettnerites. Paulinites has also been

used by, among others, Martinsson 1 960; Kielan:Jawor

owska 1966; Szaniawski 1 970; on species from Europe.

However, in my opinion none of the European species

belong to Paulinites.

The following scolecodonts from the Palaeozoic of

North America might be placed in the genus Kettnerites (I

have not noted any species in com mon between the North American and Baltic faunas) : O Nereidavus Grinnell 1 877

(N. ontarioensis Stauffer 1 939, p. 507, Pls. 57 : 1 2 , 13, 15 ;

58 :26, 32, 34; N. planus Stauffer 1939, pp.507-508, P l . 57:9,

1 0; N. invisibilis Eller 1 940, pp. 1 6--17 , Pl. 2 : 1-1 1 ; N. harbin

sonae Eller 1941 , pp. 325-326, Pl. 37: 1 , 2, 4, 5; N. angulosus

Eller 1 945, pp. 1 90-1 9 1 , Pl. 7 : 1 2-16; N. exploratus Eller

1 963a, p. 1 77, Pl. 1 :20, 2 1 ; N. incrassatus Eller 1 964, p. 231 ,

Pl. 1 : 7-13 ) . O Arabellites Hinde 1 879 (A. arcuatus Hinde

1 879 - Stauffer 1 939, p. 501 , Pl. 58: 1 7; A. priscus Stauffer

1 939, p. 504, Pl. 58 : 1 8; A. dauphinensis Stauffer 1 939, p. 502,

Pl. 58:4; A. sinuatus Walliser 1 960, pp. 22-23, Pl. 5 :3a-c) .

O Oenonites Hinde 1 879 ( O. bidens Eller 1 940, pp. 26--27, Pl. 5 :3-5) . OIldraites Eller 1 936 (J. geminus Eller 1 940, pp.

29-30, Pl. 6 : 1-5; l. eminulus Eller 1 963a, p. 1 76, Pl. 1 : 1 8) .


OPolychaetaspis Kozlowski 1956 (P. ? kozlowskii Walliser 1 960,

pp. 26--27, Pl. 6 : 1-3) .

Type species. - Kettnerites kozoviensis Zebera 1 935.

Subgeneric taxonomy. - Inside the genus Kettnerites two sub

speeies, Kettnerites ( Kettnerites) Zebera 1 935 and Kettnerites

(Aeolus) Bergman 1987 can be recognized on basis of the

following features: The MI 's of K. (Aeolus) are fairly slender

jaws of ten with a fairly wide basal portion and on the

slender MIl ' s the pre-cuspidal dentieles, if any, are small .

Kettnerites ( K. ) abraham Bergman 1 987 Figs. I 3N-P, I SF, 26 , 27,

Synonymy. - 0 1 987 Kettnerites abraham n .sp. - Bergman, p.

45 , Figs. 1 3N-P, 18, 26, 27.

Derivation of name. - As for the nominal subspecies.

Holotype. - As for the nominal subspecies.

Type locality. - As for the nominal subspecies.

Type stratum. - As for the nominal subspecies.

Material. - Fig. 2 ; more than 241 right MI, 205 left MI, 309

right MIl, and 295 left MIl .

Occurrence. - Figs. 4 and 5; Late Llandovery to Early Wen

lock. Lower Visby Beds, unit b, to H6gklint Beds unit d and

southwest facies.

Diagnosis. - Right MI: Dentieles normal to large size, slightly

slanting, covering inner margin from fakal arch to un

denticulated ridge . Shank ends in a sharp tip; almost

rounded, large, thin-walled flange, with a spur or a spur

like structure .

Left MI: Dentieles of normal to large size, covering inner

margin from fakal arch to the undenticulated ridge . Al

most rectangular inner wing. Spur or spur-like structure

large to minute .

Right MIl : Two large pre-cuspidal dentieles of equal size,

followed by a large eus p and a smaller dentiele . Shank

subparallel; ramus short and wide.

Left MIl: Double cusp, parts arranged laterally almost

side by side. Outer, slightly more anterior cusp, somewhat smaller. Intermediate dentary pronounced.

Description. - Right MI, dorsal side: Length 0 .34-1 .51 mm, width about lA of the jaw length. The jaw is fairly rounded

in cross-section. The inner margin increases in convexity

from K. (K.) abraham abraham to K. ( K. ) abraham isaac. The

dentary covers the total length between the fakal arch and

the undenticulated ridge with moderate, equal-sized denti

eles in the nominal subspecies and larger unequal-sized

dentieles in the younger subspecies. The basal portion has

a large flange with a distinet spur on the nominal subspe

eies and it becomes very conspicuously shaped and large

but with a minor spur-like structure in K. (K.) abraham

isaac. The shank ends sharply.

Left MI, dorsal side : Length 0.33-1 .80 mm, width some

what less than lA of the jaw length . The inner margin is

slightly convex on the nominal subspecies and becomes



s-shaped on K. ( K. ) abraham isaac. The denticles vary from

moderate in size to large , covering the inner margin from

the fakal arch to the undenticulated ridge. The inner wing

is almost rectangular. In the nominal subspecies, the basal

portion is fairly large with a prominent spur; in K. ( K. )

abraham isaac i t becomes very large with a conspicuous

outer side, but with a minor spur-like structure.

Right MIl, dorsal side: Length 0 .26-1 .41 mm, width

about half the jaw length . Two large pre-cuspidal denticles

of equal size are present, with the anteriormost to the right

of the posterior one. The cusp is large, slanting posteriorly,

followed by one minor denticle . The denticulated ridge is

composed of9-l0 large denticles, which slant and decrease

in size towards the posterior. The shank occupies about

half the jaw length, tapers slightly to the posterior and

terminates in a rounded posterior extremity. The bight is

shallow; the bight angle varies from acute to a right angle.

The ramus is short and wide. The anterior outer margin is

almost straight or has a small concavity.


half the jaw length or slightly more . The parts of the double

cusp are positioned almost side by side , the outermost

being slightly smaller. The intermediate dentary is pro

nounced, composed of 4-7 denticles of equal size. Poste

rior to these, on the denticulated ridge, around nine fairly

large denticles slant towards the posterior. The shank ta

pers slightly to the posterior, occupying about half of the

jaw length. The inner wing is alm ost triangularly rounded,

representing about half the jaw length . The bight is shal

low. The ramus is almost triangular, fairly wide, with a

convexity occupying the main part of the anterior margin .

Discussion. - The most characteristic feature to differentiate

the two subspecies from each other is the shape of the basal

portion. The long s-shaped outer margin of the basal por

tion of K. (K.) abraham isaac is very conspicuous.

Fig. 26. Kettnerites (K) alYraham alYraham All speeimens except N and O are in dorsal view, x 120. DA. Left MI, LO 5364: 1 , Buske l , Lower Visby Beds, unit e , 79-4lLJ. D B. Right MI, LO 5363:2, Lickershamn 2, Lower Visby Beds, unit f, 73-53LJ. D e. Left MI, LO 5365: 1 , Nygårdsbåckprofilen l, Lower Visby Beds, unit e, 79-42LJ. O D. Left MI, LO 5364:2, same sample as A. D E. Right MI, LO 5363:3, same sample as B. O F. Left MI, left MIl and MIll fused together, LO 5360:9, Sjåiso l, Lower Visby Beds, unit b, 79-l 2LJ. O G. Holotype, right MI, LO 5364:3, same sample as A. OH. Right MI, LO 5362:4, Buske l, Lower Visby Beds, unit e , 79-40LJ. Ol . Right MIl, LO 5367:5, Håftingsklint l , Upper Visby Beds, 76-9CB. OJ. Left MIl, LO 5760: 1 , Vattenfallsprofilen l, Upper Visby Beds, lowermost part, 70-l 4LJ. O K. Right MIl, LO 5368: l, Vattenfallspnr filen l, Up per Visby Beds, 76-8LJ. OL. Left MI, LO 5363:4, same sample as B. OM. LeEt Ml, LO 5362:5, same sample as H. O N. Left MI, LO 5365:2 , same sample as C, x80. 00. LeEt MI, LO 5368:2, same sample as K, x80. O P. Right MI, LO 5360: 1 0, same sample as F.


Kettnerites ( K. ) abraham abraham Bergman 1 987 Figs. l 3N, o, l 8F, 26 , 27B, D, E, I ,]

Synonymy. - Kettnerites (K.) abraham abraham n. subsp. - Berg

man, pp. 47-49, Figs. l 3N, 0, 18 , 26, 27B, D, E, I , j.

Derivation of name. - In honour of my son, Carl Abraham.

Holotype. - LO 5364:3 , right MI, Fig 26G.

Type locality. - Buske 1 .

Type stratum. - Lower Visby Beds, unit e .

Material. - Fig. 2 ; more than 1 79 right MI, 1 46 left MI, 202

right MIl, and 197 1eft MIl .

Occurrence. - Figs. 4 and 5; Late Llandovery to Early Wen

lock. Lower Visby Beds, unit b, to Hogklint Beds unit b .

Buske l , Gnisvard l and 2 , Gustavsvik l , 2 , and 3 , Hiiftings

klint l and 4, Halls Huk l , Ireviken l , 2, and 3, Korpklint

l , Lickers l , Lickershamn 2 , Nygårdsbiickprofilen l , Ny

hamn l , 2, and 4, Ronnklint l , Saxriv l , Sjiilso l , Sniick

giirdsbaden l , Vattenfallsprofilen l , Ygne 2 .

Diagnosis. - Right MI : Large, slightly slanting denticles of

almost equal size, covering inner margin. Shank ending

with sharp tip; subrounded large thin-walled flange, pro

nounced spur. In dorsal view the myocoele opening is

tubular.

Left MI: Denticles large, decreasing in size toward poste

rior end. Undenticulated ridge tapering posteriorly, pro

nounced spur.

Right MIl : Two large, pre-cuspidal denticles of equal

size, followed by a large cusp and a smaller denticle . Shank

almost parallei, ramus short and wide.

Left MIl: Double cusp, parts arranged laterally almost

side by side, the outer, slightly more anterior cusp is some

what smaller. Intermediate dentary pronounced.

Description. - Right MI, dorsal side : Length 0 .34-1 .51 mm,

width about 1/4 of length . The inner and outer margins are

almost parallei, in some cases slightly tapering. The ante

rior end forms a fairly large fang. The coarse denticulation

extends along the convex inner margin for about 0 .7 of the

jaw length . About 20 denticles, slanting except for the anteriormost ones, and almost equal in size, end as one or

a few small knobs immediately anterior to the inner wing.

The transition between the inner margin and the undentic

ulated ridge is at a lateral , outward bend of the ridge . The 'undenticulated ridge is fairly high, narrow, forming the posteriormost, sharp tip of the shank. The inner wing is of

normal size, representing about a quarter of the jaw length .

The inner wing is deeply downfolded, almost triangular; its

anterior part wide and rounded. The basal furrow is wide

and long.

The basal plate is not recorded.

The flange is thin-walled, large , subrounded, slightly

down bent along its outer margin and its anterior part

elevated. The flange en ds in a small concavity, anterior of

which is a spur, representing the outermost part of the

ligament rim. The basal portion is short and in dorsal view

seems to end like a tube. The basal angle is usually about



25-30° . The outer margin anterior to the basal portion has

a slight concavity in the middle part.

Ventrai side : The strongly enclosed myocoele opening

represents about V3 of the jaw length . A shallow furrow

surrounds the flat myocoele rim along its anterior part.

The rim is elevated, forming two high, short, narrow and

sharp triangular ridges on the anterior inner and outer

margin . The spur is distinct. A sharp ridge continues along

the inner margin of the shank.

Left MI, dorsal side: Length 0.34-1 .80 mm, width less

than lf4 of length. The inner and outer margins are almost

parallei, ending anteriorly in a normally developed fang.

The inner margin is slightly convex to almost straight with

about 20 large ( the anteriormost on es of almost equal size) ,

slightly spaced denticles ending just anterior to the inner

wing. The transition from the inner margin to the un

denticulated ridge is smooth . The undenticulated ridge is

low and becomes narrower toward the posterior end,

where it alm ost reaches the posterior margin . The inner

wing is of normal size, slightly more than 0.2 of the jaw

length, almost rectangular and slightly downfolded. The

basal furrow is of normal length, deep and parallei to the

outer margin of the basal portion, which has a downfolded

posterior part. On the anteriormost outer part of the basal

portion there is a concavity and a spur similar to the one on

the right MI.

Ventrai side: The strongly enclosed myocoele opening

represents about one third of the jaw length . The ligament

rim is narrow and somewhat depressed in relation to the

ventrai surface . Along the inner margin of the opening and

the inner wing, a narrow, fairly low ridge is present. On the

outer posterior part there is a high, narrow, sharp ridge

forming a sp ur. The spur dominates the small low ridge

which con tinues along the whole of the outer margin of the

basal portion.

Right MIl, dorsal side : Length 0 .26--1 .35 mm, width

about half the length . Two large pre-cuspidal denticles of

equal sized, with the anteriormost one positioned to the

right of the posterior one . The cusp is large , slanting

towards the posterior, followed by one minor denticle. The

denticulated ridge is laterally convex, composed of 9-1 0

large denticles, which decrease in size and slant towards the

Fig. 2 7. A, C, F, G, H, K: Kettnerites (K.) abraham isaac; B, D, E, I , ]: K. (K.) abraham cf. abraham All specimens in dorsal view. DA. Left MIl, LO 5767:5, Vattenfallsprofilen l , H6gkJint Beds, unit b, 70-6LJ, x1 20. DB. Right MIl, LO 5764:6, Vattenfallsprofilen l , H6gklint Beds, unit a, 70-20LJ, x 1 20. OC. Left MIl, LO 5772:5, Stave l, Slite Beds, Slite Mari, central part, 75-1 1 CB, x1 20. OD. Right MIl, LO 5368:3, Vattenfallsprofilen l , Upper Visby Beds, 76-8LJ, x80. DE. Left MIl, LO 5765: 1 , Lickershamn 2, H6gklint Beds, unit a, 73-57LJ, x 1 20. OF. Vattenfallsprofilen l, H6gklint Beds, unit d, Valdana testudo level, sample RM; Fl left MI, AN 271 3, x80; F2 right MI, AN 271 4, x80. OG. Vattenfallsprofilen l , H6gkJint Beds, unit b , 70-6LJ; Gl , holotype, left MI, LO 5767:7, x80; G2 right MI, LO 5767:6, x80. OH. Right MIl, LO 5763:4, Ansarve l, H6gklint Beds, SW facies, upper part, 79-46LJ, x80. DI . Right MIl, LO 5760:2 , Vattenfallsprofilen l , Upper Visby Beds, 70-l 4LJ, x1 20. 0J. Right MI, LO 5764:7, same sample as B, x120.

D K. Same sample as F; Kl left MI, AN 2715 , x80; K2 right MI, AN 2716, x80 .


posterior. The shank, occupying about half the jaw length,

tapers slightly to the posterior, with a rounded posterior

extremity. The posterior outer margin is almost straight to

slightly concave . The bight is shallow; the bight angle varies

from about 55° to 75° . The ramus is short and wide. The anterior outer margin is usually nearly straight, though a

small sinus may exist.


represents �3_3/4 of the jaw length . The anterior and inn er

margins are slightly curved and surrounded by a fairly thin

rim.

Left MIl, dorsal side : Length 0 .38 mm-l .48 mm, width

about half the length or slightly more. The double cusp

with its slightly smaller outer part positioned somewhat

posteriorly, points almost at right angle to the extension of

the denticulated ridge or has a slight posterior slant. The

intermediate dentary is pronounced, composed of 4-7

denticles irrespective of the size of the jaw. The denticles

are of equal size, except the posteriormost one, which is

normally larger. The 8-1 0 fairly large denticles of the

denticulated ridge slant posteriorly and are of alm ost equal

size or decrease slightly towards the posterior. The shank

occupies about half of the jaw length and tapers slightly to

the posterior. The inner wing is about half of the jaw length

and almost triangular, rounded. The bight is shallow; the

bight angle varies from acute to right. The ram us is nearly

triangular, fairly wide, with a convexity occupying the main

part of the anterior margin.


represents V2-% of the jaw length . The almost straight

anterior and inn er margins of the myocoele are sur

rounded by a rim of normal width .

Discussion. - Two different dentary forms among the MI's

have be en recorded, viz . the usual form with large denticles

(e .g. Fig. 260, P) and a less common form with fairly thin

and more numerous denticles (e .g. Fig. 26D, E) .

Comparison. - The right MI ( 738/5 1 ) reported as Kettnerites

polonensis by Mannil & Zaslavskaya ( l 985a) is very similar

(conspecific?) to K. ( K. ) abraham abraham (see chapter

'Comparison with paulinitid faunas from other areas ' ) .

K.ettnerites ( K. ) abraham isaac Bergman 1 987 Figs. 1 3P, 1 8F, 27A, c, F-H, K

Synonymy. - D 1987 Kettnerites (K. ) abraham isaac n.subsp . -

Bergman, pp. 49-51 , Figs. 1 3P, 18 , 27A, C, F-H, K

Derivation ofname. - Isaac, on the Abraham ' lineage ' .

Holotype. - L O 5767:7, left M I , Fig. 27G: 1 .

Type locality. - Vattenfallsprofilen 1 .

Type stratum. - Hogklint Beds, unit d , Pterygotus Beds.

Material. - Fig. 2 ; more than 62 right MI, 59 left MI, 1 07


Occurrence. - Figs. 4 and 5 ; Early Wenlock. Hogklint Beds,

unit b, c , d, and the SW facies. Ansarve l, Ar l, Lauter l ,

Langhammarsviken 2 , Lauterhornsvik 3 , Lickershamn 2,

54 Claes F. Bergman

Nors Stenbrott l , Saxriv l , Stutsviken l , Svarven l , Vatten

fallsprofilen l .

Diagnosis. - Right MI: Denticles increase in size from nor

mal on fakal arch to large on posterior part of the dentary

along the reverse s-shaped inner margin . Large, thin-walled

flange, long, s-shaped outer margin of basal portion, end

ing in a spur-like structure . Shank ends sharply.

Left MI: Anterior taper, inner margin convex, covered by

fairly large denticles. Outer margin of basal portion rela

tively long.

Right MIl : Two large pre-cuspidal denticles of equal

sized, followed by a large cusp and a smaller denticle .

Shank subparallel; ramus short and wide.

Left MIl: Double cusp, parts arranged almost side by

side. The outer, slightly more anterior cusp, is somewhat

smaller. Intermediate dentary pronounced.


width 1f4 of length. The jaw tapers anteriorly from the fairly

wide basal portion, ending in a normally developed fang,

bent slightly upwards in relation to the dorsal surface . The

convex inner margin has 25-30 fairly large to large denti

cles from the fakal arch posteriorly, ending abruptly just

before the inner wing. They slant and increase in size

posteriorly, with the exception of the somewhat larger

anteriormost one and the slightly smaller posteriormost

ones. The undenticulated ridge continues as a smooth

ridge, slightly bent towards the inner margin, forming the

posteriormost tip of the sharply pointed shank. The inner

wing is of normal size, downfolded, alm o st triangular, with

a rounded prominent anterior ' corner' . The outer margin

of the shank is sigmoidal; the basal furrow is wide and long.


The flange is large, thin-walled, e!evated, the margin

s-shaped and rounded, ending in a minor concavity and a

spur-like structure (Figs. 27G:2 , 27K:2 ) at the anterior part

of the basal portion, where the ligament rim meets the

flange . The posterior, outermost part of the flange is moderately downfolded. The basal portion is wide and fairly

long. The basal angle is about 35° . The outer margin, anterior to the basal portion, is concave, ending in the sickle-shaped falx.

VentraI side : The strongly enclosed myocoele opening

represents about one third of the jaw length . The anterior part of the opening is surrounded by a flat ridge which in

turn is surrounded by a narrow groove . The rim increases

in size toward the posterior inner and outer margins. It

forms a ridge in the posteriormost inner margin of the

shank.

Left MI, dorsal side: Length 0.33-1 .37 mm, width slightly

less than 1f4 of length. The jaw tapers from basal portion to

the pointed fang, which is moderately bent upwards in

relation to the dorsal surface . The inner margin is convex,

denticulated along its entire length, representing about 0 .7

of the jaw length, extending from the fakal arch posteriorly

and reaching the inner wing. The denticles are fairly large,

of equal size, slanting, and slightly spaced. The anterior

most 3-4 denticles are of ten the largest and point forward.

The undenticulated ridge forms a convex continuation of


the denticulated inner margin, starting beside the anterior

most part of the almost rectangular, normal-sized inner

wing. The inner margin of the wing is nearly paralleI to the

undenticulated ridge . The basal furrow is short and shal

low, oriented intermediately between the undenticulated

ridge and the outer margin of the basal portion . The basal

angle varies from 25' to 40' . The outer margin of the basal

portion is long, downfolded along the posterior half, form

ing a small outer wing on the anterior part and ending in a

concavity. Posterior to the concavity is a spur-like extension

of the ligament rim. The outer margin is concave with the

sickle-shaped falx occupying the anterior half.

VentraI side : The strongly enclosed myocoele opening

represents about V3 of the jaw length . It is elongated,

surrounded by the ligament rim, the surface ofwhich is flat

to concave on its anterior part but forms a high, narrow

ridge along the inner margin . On the anterior, outer mar

gin of the basal portion, a small ridge with a furrow along

its outer margin grows into a narrow, relatively high ridge

on the posteriormost part.

Right MIl: Very close resemblance to the right MIl of K.

( K. ) abraham abraham. Among some K. ( K. ) abraham isaac

specimens, the sinus in the anterior outer margin is more

pronounced, and the shank is slightly less tapering. Length

0.26--l .41 mm.

Ventrai side : The slightly enclosed myocoele opening is

very similar to the nominal species, but the anteriormost

inner part of the rim has a protruding corner.

Left MIl: Very close resemblance to the left MIl of K. ( K. )

abraham. The pre-cuspidal denticle seems to b e more nee

dle-shaped among the K. ( K. ) abraham isaac than in the left

MIl of K. ( K. ) abraham abraham. Length 0.33-l .00 mm.

VentraI side : Very similar to the nominal species.

Remarks. - At present the MIl is not so well preserved, and

the material is fairly small at each locality. The variability of

the morphological characteristics of the MIl of the two

subspecies overlaps. Although some differences noted in

the description are at hand, the morphological characters

overlap and are thus not useful for identification. It is not improbable that with a very large, well preserved material ,

some other characteristics for distinguishing the MIl of the two subspecies will be found. More probably, however,

additional material will increase the amount of overlap and

thus continue to make the identification of the MIl on the

subspecific leve! further impossible.

Discussion. - The transition from Kettnerites (K.) abraham

abraham to K. (K.) abraham isaac is observable in the Hog

klint Beds, unit b, e .g . Vattenfallsprofilen l ( 70-6LJ) and in

Lickershamn 2 (73-57LJ, 73-65LJ, 75-1 05CB samples not

included in the locality list) .

Kettnerites ( K ) jacobi Bergman 1 987 Figs. I 3Q, I SF, 2SA, B

Synonymy. - 0 1 987 Kettnerites ( K. ) jacobi n.sp. - Bergman,

pp. 51-53, Figs. 1 3Q, 1 8 , 28A, B .

Derivation ofname. - In honour of my son Jakob.


Fig. 28. Kettnerites (K.) jacobi. All specimens in dorsal view. DA. Slitebrottet 2, Slite Beds, Slite Mari, 83-3I LJ; Al left MI, LO 5773:5, x80; A2, holotype, right MI, LO 5773:6, x120; A3 1eft MIl, LO 5805:2 , x1 20; A4, right MIl, LO 5805: 1 , x 120. DB. Stutsviken l , Hogklint Beds, unit c , 77-7CB; Bl left MI, LO 5769: 1 , x80; B2 right MI, LO 5769:2 , x80 .

Holotype. - LO 5773:6, right MI, Fig. 28A:2 .

Type locality. - Slitebrottet 2 .

Type stratum. - Slite Beds, unit g.

Material. - Fig. 2 ; More than 30 right MI, 27 1eft MI, 39 right

MIl, and 37 left MIl .

Occurrence. - Figs. 4 and 5; Wenlock. Slite Beds, unit f, Slite

Mari, unit g, unit Lerberget Mari - Pentamerus gothlandicus;

Broa 2, Haganås l , Myrsne l , Slitebrottet l and 2, Stave l , Stora Mafrids 2 , Tjeldersholm l , Val bytte l , Vallstena 2 .

Diagnosis. - Right MI : Large jaws with 1 5-20 large , anteri

orly widely spaced denticIes, decreasing in size and becom

ing more densely spaced toward the posterior end. Shank

56 Claes F. Bergman

sharp-ended; flange thin-walled, rounded, with spur-like

structure.

Left MI: Almost a mirror image of the right MI except for

the basal portion. Inner wing anteriorly rounded, spur-like

structure distinet.

Right MIl: Two large pre-cuspidal denticles; cusp of mod

erate size; large wide shank, ending fairly pointed.

Left MIl: Thin needle-shaped pre-cuspidal denticle;

large cusp; pronounced, slightly spaced intermediate den

tary; subtriangular ram us.

Description. - Right MI, dorsal side : Length 0.55-1 .68 mm,

width about V3 of length . The surface is flat anteriorly and

convex posteriorly. The jaw tapers slightly anteriorly, end

ing in a well developed fang, bent some 10 ° upwards in

re!ation to the surface . Slightly convex to almost straight

inner margin. The large, 1 5-20, normally 1 6--18 denticles,

anteriorly slightly spaeed, decrease in size toward the pos

terior end and are slightly slanting or perpendicular to

wards the extension of the inner margin, covering about

0.6 of the total jaw length . The denticulated ridge ends in

small denticles and continues as the undenticulated ridge

well before reaching the same leve! as the an teriormost part

of the inner wing, or on the outer side , before reaching the

bight. The undenticulated ridge is smooth, high and long,

representing 0.28-0.35 of the jaw length. It is almost

straight, sometimes slightly bent towards the outer margin,

forming the posteriormost part of the jaw and ending in the

sharp tip of the shank. The inner wing is of normal size,

about 1/4 or less of the jaw length, almost triangular, with

rounded corners, the widest part on the anterior half,

steeply downfolded.


The basal furrow is short, squeezed between the un

denticulated ridge and the large, rounded, elevated, thin

walled flange . The posterior, outermost part of the flange

is downfolded. Anterior to the flange there is a small con

cavity in connection with a vague rim and a spur-like exten

sion (Fig. 28A:2) . The basal portion is of normal size, alm ost triangular with a basal angle of about 30-40° . The outer margin anterior to the basal portion is rounded with

a shallow concavity in the middle, ending anteriorly in a

sickle-shaped falx.

VentraI side : The strongly enclosed myocoe!e opening

represents about 1/3 of the total jaw length . The ligament

rim surrounds the opening along the anterior and inner

margins and forms a small spur at the anterior, outer

position .

Left MI , dorsal side : Length 0 .5 1-2.40 mm, width 1/4 of

length . The dorsal surface is slightly concave. The jaw

tapers gradually towards the anterior, ending in a pointed

fang, moderately bent upwards in relation to the surface .

The inn er margin is faintly to moderately convex, covered

for about 0.6 of the jaw length by 15-20 (normally 1 6--18)

denticles, of which the anteriormost 5-6 are very large and

widely spaeed; the denticulation ends well before reaching

the leve! of the inner wing. The denticles slant slightly,

decreasing in size posteriorly, ending as small knobs. The

anteriormost denticle is sometimes slightly smaller than the

second and of ten directed forward. This is occasionally also


the case with the anterior second and third denticles.

There is a smooth transition to the long, straight, low and

narrow undenticulated ridge . The inner wing is of normal

size, about 1/4 of the jaw length, rounded, moderately down

folded, forming the posteriormost part of the shank. The

basal furrow is short, narrow and oriented slightly toward

the outer margin. The posterior half of the outer margin of

the basal portion is somewhat downfolded; the anterior

half has a small concavity with a spur-like rim in front of it.

The basal portion is of normal size, skew, with a basal angle

of about 30-40° . In the middle of the outer margin, at its

transition into the fang, and anterior to the basal portion,

the re is a vague concavity.

VentraI side : The strongly enclosed myocoele opening,

V4-V3 of the jaw length, is in its slightly curved anterior part

surrounded by a low ligament rim. Along the outer margin

the rim becomes more distinet with anterior spur-like ex

tensions.

Right MIl, dorsal side : Length 0 .35-2 . 1 2 mm, width

more than half the jaw length . The inner margin is slightly

convex. Two large pre-cuspidal denticles form the

rounded, almost pointed anterior margin. Together with

the cusp they are bent strongly upwards and slant slightly

towards the posterior. The cusp is of moderate size, fol

lowed by a minor intermediate denticle. The post-cuspidal

dentary comprises 8-9 large denticles, decreasing in size

and slanting slightly towards the posterior. The shank,

more than half the jaw 1ength, is wide with almost paralleI

sides. The bight is fairly deep, the bight angle about 50-60° .

The ramus i s large , almost triangular, with an extremity

pointing slightly posteriorly. The anterior outer margin

varies from slightly convex to concave .


represents %, or somewhat less, of the jaw length . The

anterior and inner margins are almost straight and sur

rounded by a fairly thin rim with a corner slightly protrud

ing anteriorly on the anterior inner margin .

Left MIl, dorsal side: Length 0.42-2 .00 mm, width about

half the jaw length . The cusp is large, with a small needle

shaped pre-cuspidal denticle along the anterior sickleshaped margin, oriented in the same direction as the cusp . The intermediate dentary has 6--8, slightly spaced denticles

of normal size. There is an abrupt transition to the post

cuspidal dentary, which consists of 7-1 0 fairly large denti

cles, slanting slightly towards the posterior. The shank,

about half the jaw length , tapers slightly towards the poste

rior. The inner wing is almost triangular, laterally slightly

less extended than the shank, its anterior part being widest.

The bight is normal; the bight angle about 70° ; the ramus

is fairly large, nearly triangular with its extremity pointed

slightly posteriorly. The anterior outer margin is smoothly

sigmoidal .


represents about half, or slightly more, of the jaw length .

The opening is smoothly curved, surrounded by a rim of

normal width .

Discussion. - The MI elements of Kettnerites ( K ) abraham

abraham, K ( K ) abraham isaac and K ( K ) jacobi are char

acterised by a large, rounded, thin-walled flange , a spur


decreasing in size from late Visby time, and large denticles

which increase in size from Visby time to Slite time. The

denticle formula of the MIl is persistent through time,

though the size of the pre-cusps decreases to a needle-like

pre-cusp in the Slite Beds. K. ( K. ) abraham abraham evolves

gradually into the Hogklint subspecies, i .e . K. (K.) abraham

isaac. It is possible that K. (K.) jacobi has evolved from the

Hogklint type (Figs. 1 3N-Q) . Supporting this theory are

the equivalent shape and size of the flange and the dentic

ulation of the left and right MI, and the similar denticle

formula of the MIl 's . However, there is a lack of informa

tion from the lower Slite Beds.

Remarks. - In the more than 14 samples yielding K. ( K. ) abraham isaac, the MIl i s consistently recorded about twice

as frequently as the MI. There are a number of conceivable

explanations for this discrepancy. First, the MIl, with its

denticle formula, is easier to determine than the MI. Sec

ond, I might have included other MIl ' s belonging to some

unidentified polychaete taxa in the count. Third, the MI is

more fragile than the MIl. Fourth, although this is difficult

to prove, the MIl may be represented by a double number

of jaws in each apparatus. Fifth, the recorded frequencies

are caused by the random variation in the preservational

processes. Most probably the skew proportion of the two

jaws is a question of preservation and identification.

Kettnerites ( K. ) bankvaetensis Bergman 1 987 Figs. 1 3I-M, 1 8E, 29-32

Synonymy. - O 1 987 Kettnerites ( K. ) bankvaetensis n.sp . - Berg

man, pp. 53-56, Figs. 1 3I-M, 1 8 , 29-32 .

Derivation of name. - Latin bankvaetensis, from the type local

ity Bankvat l , where this species is common and dominates

the annelid fauna in some leveis.

Holotype. - LO 5829:5 , right MIl, Fig. 31F.

Type locality. - Bankvat l .

Type stratum. - Hamra Beds, unit b .

Material. - Figs . 2 and 3; more than 250 elements each of

right and left MI ' s and MIl 's .

Occurrence. - Figs. 4 and 7; Early Wenlock to Late Ludlow,

Hogklint to Hamra Beds, unit c, Sundre Beds lower (? ) part; Angmans 2, Baju l , Bankvat l , Bofride l , Bottarve l and 2 ,

Bringes 3 , Dapps l , Faludden 2 , Fie 3, Fjale 2 and 3 ,

Fjardinge l , Gane 2 , Gannor l , Garnudden 3 and 4, Gisle

l , Gothemshammar 2 and 6, Gutenviks l , Haganas l , Hide

1 , Juves 4, Kiittelviken 5, Kauparve l , Krakfot l , Kroken 2 ,

Kullands 2 , Mollbos l , Nabban 2 , Nyan l , Nygårds 2 , Ollajvs

l , Slitebrottet l , Stora Myre l , Storugns lB , Strands l ,

Suderbys 3 , Valby Bodar l , Valleviken l , Vallstena 2 , Vaster

bjars l , Vike 2 and 3, Yxne 1 .

Diagnosis. - Right MI: Inner margin almost straight with

closely spaced denticles of normal size. Shank sharply

ended, flange angular, highly elevated posteriorly.

Left MI: Inner margin almost straight, denticles decrease

in size posteriorly, ending without reaching the practically


rhombic inner wing. Ligament rim forms a short ridge on

posteriormost outer part of basal portion .

Right MIl : Two fairly large pre-cuspidal denticles, the

anterior one slightly larger; large cusp followed by one or

two fairly small intermediate denticles.

Left MIl: Double cusp with cusps of equal size, the ante

rior one positioned slightly more exteriorly. Shank tapers

to a fairly pointed extremity.


width about lf4 of length. The inner and outer margins

taper slightly toward the anterior end: a normally devel

oped fang, moderately bent upward in relation to the

dorsal surface . The inner margin is slightly convex among

Wenlock jaws while it is straight among the Ludlow ones.

The inner margin is covered by 24-44 normal-sized, close

spaced denticles, which are more or less perpendicular to

the extension of the inner margin . The denticles occupy

about 0 .6-0. 7 of the jaw length, and the number of denti

cles is to some degree coupled to jaw length . They extend

from the anteriormost part of the falcal arch down to the

inner wing, except in specimens from the Wenlock, where

the denticulation ends before reaching the inn er wing.

The denticles are almost equal in size except for the poster

iormost ones which decrease in size, ending as crenula

tions. The transition between the denticulated inner mar

gin and the undenticulated ridge is characterized by

crenulations on a small, winding, narrow ridge on top of

the main ridge . The undenticulated ridge is fairly narrow,

high, and its anterior part bends toward the outer margin .

It continues down to the posteriormost part of the jaw,

forming the sharp tip of the shank. The inner wing is of

normal size, about a quarter of the jaw length, rounded and

deeply downfolded, with its widest part at the anterior end.


The basal furrow is fairly long and deep posteriorly. The

flange is angular, the posterior part highly elevated, with

the posteriormost part somewhat downfolded, and the an

terior part abutting the anterior end of the jaw. The liga-

Fig. 29 (p . 58) . Kettnerites (K.) bankvaetensis. All specimens in dorsaJ view, x 1 20. OA. Right MI, LO 5836:7, Gandarve l, Halla Beds, 71-8 ILJ. OB. Gothemshammar 6, Halla Beds, unit c, 73-24LJ; B l left M I , L O 5795: l ; B2 right MI, L O 5795:2 ; B3 left MIl, L O 5795:3 ; B4 right MIl, LO 5795:4. O C. Vike 2, Slite Beds, Pentamerus gothtandicus Beds, 83-4CB; Cl left MI, LO 5782: 1 ; C2 right MI, LO 5782:2 ; C3 leEt MIl , LO 5782:3 ; C4 right MIl, LO 5782:4 . OD. LeEt MI, LO 5836:6, Gandarve l , Halla Beds, 7 1 -8ILJ. O E . Gothemshammar 2, Halla Beds, unit c, 75-35CB; El leEt MI, LO 579 1 : 1 ; E 2 right MI, L O 5791 :2 ; E 3 leEt MIl, L O 579 1 :3 ; E 4 right MIl,

LO 5791 : 4.

Fig. 30 (p. 59) . Kettnerites (K.) bankvaetensis. All specimens in dorsal view. O A. Mollbos l, Halla Beds, unit b , 77-28LJ; Al leEt MI, LO 5792: 1 , x60 ; A2 right MI, LO 5792:2 , x60; A3 leEt MIl,

LO 5792:3, x60; A4 right MIl, LO 5792:4, x60. O B. Gothemshammar 6, Klinteberg Beds, unit a, 73-30LJ; Bl leEt MI, LO 5799: 1 , x80; B 2 right MI, LO 5799:2 , x80; B 3 left MIl, L O 5799:4, x80; B4 right MIl, LO 5799:5, x1 20; B5 left MIl, LO 5799:6, x1 20; B6 leEt MIl, LO 5799 :3, x80. OC. Fjardinge l , Klinteberg Beds, unit b, 77-5CB; Cl left MI, LO 5796: 1 , x80; C2 right MI, LO 5796: 2, x80; C3 right MIl, LO 5796:3, x80.


Fig. 29 (caption on p. 57) .

FOSSILS AND STRATA 25 ( 1 989) Silurian paulinitid polychaetes from Gotland 59




ment rim forms a short ridge with a spur-like structure

along the anteriormost outer margin of the basal portion

(Figs. 30A:2 , B :2 , C:2 , 3 1D , H) . The basal angle is between

28° and 50° .

Ventrai side : The strongly enclosed, rounded myocoele

opening, represents about 1f4 of the jaw length . The curved

opening is surrounded by a broad and flat ligament rim,

which rises to a fairly small and narrow ridge on the ante

rior outer part of the inner wing.

Left MI, dorsal side: Length 0.46-1 .75 mm, width about 1f4-lfs of length. The inn er and outer margins are almost

parallei, ending anteriorly in a normally developed fang

which is slightly bent upward compared with the relatively

flat dorsal surface . The inner margin is slightly convex

among the Wenlock jaws but almost straight among the

Ludlow ones. The inner margin is covered by 30-48 closely

spaced denticles of normal size, extending posteriorly from

the fakal arch to well in front of the inner wing. The

denticles decrease in size toward the posterior, where they

end as small knobs. There is a smooth transition to the

undenticulated ridge, which continues as a low, narrow

and straight ridge down to the posterior margin. The inner

wing is of normal size, almost rhombic, somewhat rounded,

with its inner margin parallei to the undenticulated ridge.

The basal furrow, to the left of the ridge, is relative ly short,

deep and oriented parallei to the outer margin of the basal

portion. The basal angle is between 20° and 35° , and the

outer margin of the outer face of the basal portion is

downfolded, its anterior part forming a small flange or a

narrow fold, the ligament rim. The outer margin, anterior

to the basal portion, is slightly concave , followed anteriorly

by a slightly angular falx.

Ventrai side : The myocoe!e opening is strongly enclosed,

slightly crescent-shaped, and about one quarter as long as

the jaw. It is anteriorly delimited by a broad, concave

ligament rim, which forms a narrow and high ridge along

the inner side of the innermost part of the inner wing.


slightly more than half the length . There are two fairly

large pre-cuspidal denticles, the anterior one somewhat

larger and situated slightly to the right of the second one.

In three specimens the pre-cuspidal dentary is composed

Fig. 31 (opposite page) . Kettnmtes (K.) bankvaetensis. Bankvat l , Hamra Beds, unit b , 82-30CB, exeept G and H. All spe eimen are in dorsal view exeept C, D, E and J (ventrai ) . DA. Left MN, LO 5829: 1 0, x1 20. DB. MIII, LO 5829:9, x 1 20. OC. Basal part left MI, LO 5829 :8, x1 20. OD. Basal part of right MI, LO 5829:7, x120 . D E. Right MIl, LO 5829:6, x 1 20. OF. Holotype, right MIl, LO 5829:5, x120. O G. Left MI, LO 5841 : 1 , Glasskår 3, Burgsvik Beds, lowermost part, 82-1 8CB, x80. OH. Right MI, LO 5841 :2 , same sample as G, x80. DI . Left MII, LO 5829:3, x1 20. OJ. Left MII, LO 5829:4, x1 20. DK. Left MI, LO 5829 : 1 , x80. DL. Right MI, LO 5829:2 , x80.

flg. 32 (right) . Kettnmtes (K.) bankvaetensis. All speeimens in aorsal view. DA. Vasterbaekar l, Sundre Beds, middle-upper part, 75-2LJ; Al left MI, LO 5838:4, x80; A2 right MI, LO 5838:5, x80. DB. Kauparve l , Hamra Beds, lower-middle part, 76-1 3CB; BI left MI , LO 5844: 1 , x60; B2 right MI, LO 5844:2, x60; B3 left MIl, LO 5844:4, x80; B4 right MIl, LO 5844:5, x60 .


62 Claes F. Bergman

of one larger denticle followed posteriorly by two small

ones. The cusp is large, its continuation is seen as a minor

swelling of the jaw. It is followed by one or two small

denticles, the intermediate dentary. The transition to the

dentary of the denticulated ridge is marked by 2-3 denti

cles which increase in size. The 6-9 large denticles of the

ridge decrease in size posteriorly and slant in the posterior

direction. The shank, occupying slightly more than half the

jaw length, has a wide anterior part. The inner margin of

the shank is convex, the outer straight, forming a fairly

pointed extremity. The bight is fairly deep with an acute

bight angle from about 50' to 75' . The ram us is of moderate

to fairly large size, and a vague sinus can be seen along the

an terior outer margin of some jaws.


represents about �3, or slightly more, of the jaw length . A

fairly wide rim forms the almost straight anterior margin of

the myocoele opening, largest on the inner side .

Left MIl, dorsal side : Length 0 .25-2 .08 mm, width half

the length or slightly more , irrespective of the size of the

jaw. The double cusp has parts of equal and moderate size ,

the two parts being almost side by side , the anterior one to

the left. The intermediate dentary is distinet with 5-9 small

to fairly small denticles, increasing in size to the posterior.

The number of intermediate denticles and denticles on the

ridge has no relation to jaw size. The transition to the

almost straight, denticulated ridge is gradual, with the nor

mally 7-9 denticles increasing in size. The shank usually

occupies about half the jaw length; its anterior part is wide,

it taper posteriorly, and its extremity is fairly pointed,

slightly bent outwards. The inner wing is prominent, length

about half the jaw length, its almost triangular anterior part

being widest and smoothly rounded. The bight is of mod

erate size; the bight angle usually between 70' and 80' . The

ramus is fairly large and the outer anterior sinus of varying

size.


represents half, or slightly more, of the jaw length . A fairly

wide ligament rim forms the anterior margin of the open

ing, nearly straight and almost perpendicular to the extension of the jaw.

Comparison. - It is almost impossible to distinguish between

the MI ' s of K ( K ) bankvaetensis and the corresponding

element of K ( K ) martinssonii from the middle Wenlock

(e .g. Vike 2 , Slite Beds, unit Pentamerus gothlandicus or

slightly older) . From this stratigraphical leve I there is no

clear character which can be used for the differentiation of

the elements. The shape of the elements evolves, and those

from only slightly younger levels (e .g . Nygårds 2 , Halla

Beds, unit b) show at least small differences: both left and

right MI of the K (K) bankvaetensis have an almost straight

inner margin, and on the MI 's the inner wing is more

nearly rectangular. The inner wing of the right MI's of K ( K ) bankvaetensis has a straigh t inner margin and lacks the

posteriormost tip of the shank characteristic of K ( K ) martinssonii. The change of morphology continues through

time, and there is no particular similarity between the two

species in the Ludlow strata. The basal portions of the MI 's

are less angular than the almost quadratic basal portions of


K ( K ) polonensis cf. polonensis and the denticles are smaller

than in that taxon .

The MII ' s have the same denticle formula as severai of

the other paulinitid species on Gotland (e .g . , Ketinerites

(K) polonensis, K (K) burgensis, K ( K. ) abraham, K ( K ) jacobi, K ( K ) huberti, including their varieties and the valle

variety of K (A. ) sisyphi) . However, the pre-cuspidal denti

cles of K ( K ) bankvaetensis with the anteriormost denticle

slightly larger than the following one, and with the tapering

and fairly sharply ending shank, makes the element distin

guishable. The left MIl is less easy to distinguish; the

anteriormost cusp is situated slightly external to the second

cusp. The shank tapers towards the posterior and is fairly

pointed at the end.

Kettnerites ( K. ) burgensis (Martinsson 1 960) Figs. 1 3H, 18B, 33G, H, I , J, K , L

Synonymy. - 0 1 960 Paulinites burgensis n.sp . - Martinsson,

pp. 3-5 , Fig. l , 1-5. 0 1 987 Kettnerites ( K ) burgensis (Mar

tinsson 1 960) - Bergman, pp. 56-59, Figs. 1 3H, 18 , 33G-L.

Holotype. - 'Paulinites' burgensis Martinsson 1 960 pp. 3-5,

Pl. 1 : 1-5, ajaw assemblage of left and right MI and MIl, and

carriers. The carriers are not sufficiently preserved to allow

any description .

Type stratum. - Hemse Beds, Hemse Marl SE part.

Type locality. - Vastlaus l .

Material. - Fig. 3 ; more than 60 right MI, 72 1eft MI, 8 3 right

MIl, 59 left MIl .

Occurrence. - Figs. 4 and 5; Middle Ludlow, Hemse Beds,

Hemse Mari NW, unit c , d , Hemse Mari SE, and Mari

uppermost part. Bodudd 3, Botvide l , Fie 3, Gerums

kanalen l , Gogs l , Gutenviks l , Kårne 3, KJasård l , Kul

lands 2 , Linviken 2, Vaktård 2, 3, 4 and 5, Vastlaus l .

Diagnosis (emended) . - Right MI: Slender jaw, coarsely den

ticulated, anteriorly with paucidentate or with slightly spaced dentary. Pointed shank, angular, fairly thick flange ,

distinet sp ur. Left MI: Slender jaw, coarsely denticulated like right MI.

Basal portion angular. luner wing small, rounded; distinet

spur.

Right MIl: Two large pre-cuspidal denticles, cusp of mod

erate size or one large pre-cusp and a eUs p of moderate size.

Post-cuspidal dentary ending well before reaching the pos

terior margin. Slender, pointed ramus.

Left MIl: Large cusp, with or without a very large denticle

immediately posterior to it. Widely spaced intermediate

dentary with fairly large denticles. Slender shank. Almost

triangular, slender, pointed ramus.

Description. - Right MI, dorsal side : Length 0 . 1 8 to more

than 2 .61 mm, width about lf4 of the length, more among

the small specimens and less among the large ones. The jaw

tapers very slightly towards the anterior, ending in a large,

sickle-shaped fang which is only very slightly bent upwards

in relation to the almost flat dorsal surface . The inner

margin is slightly convex to almost straight. Its denticula-


Fig. 33. Kettnerites (K.) burgensis. All speeimens in dorsal view. All specimens x 120 except I , K and L. A-F: K. (K.) cf. burgensis, Vaktård 4, Hemse Beds, Hemse Mari, SE part, 81 -35LJ) . G-L: K. (K.) burgensis, Vastlaus l, Hemse Beds, Hemse Mari, SE part, 82-31LJ. DA. Left MIl, LO 5822:5. DB. Right MIl, LO 5822:6. De. Left Mil, LO 5820:3, x1 70. OD. Left MI, LO 5822:7. D E. Right MI, LO 5822:8. OF. Right MI, LO 5822:9. O G. Right Mil, LO 5806:8. OH. Left Mil, 5806:7. DI. Left MI, 5806:5, x80. 0J. Left MI, 5846:2. DK Right MI, 5846: 1 , x60. D L. Left MI,

5806:6, x80.

64 Claes F. Bergman

tion extends about :V3 of the jaw length , from the anterior

part of the falcal arch to near the inner wing. The denticles

are large anteriorly, decrease in size towards the posterior,

ending as small knobs. There are two main types of dentic

ulation as well as intermediate varieties: the extremely

paucidentate form and the form with only widely spaced

denticles. Paucidentation seems to be slightly more com

mon among the larger specimens and is almost lacking

among the smallest. Paucidentate forms have more triangu

lar denticles, laterally elongated along the length of the jaw.

The anterior denticles are usually perpendicular to the

extension of dentary, while the posterior denticles may or

may not slant slightly towards the posterior. The num ber of

denticles varies between 1 4 and 1 8, with an ave rage of 1 5 ,

and a slight positive correlation to jaw size. The undentic

ulated ridge is fairly low, short, about 0 .2 of the jaw length,

almost straight, its posterior part bent slightly to the out

side. The transition to the undenticulated ridge is very

smooth . The inner wing is almost triangular, small and

deeply downfolded, widest anteriorly with its corner vary

ing from angled to rounded. The posteriormost part of the

inner wing forms, together with the undenticulated ridge,

the sharp-ended posterior part of the fairly small, alm ost

triangular shank. The basal furrow is fairly short and shal

low.


The flange is moderately thick-walled, its posterior part

highly elevated and only slightly downfolded along the

outer posterior margin . A conspicuous spur marks the

anterior margin of the basal portion. The basal angle is

about 20-25 0 . The outer margin of the basal portion varies,

but there is always a concavity posterior to the large con

spicuous spur. The outer margin, anterior to the basal

portion, is almost straight with a wide minor concavity on

the posterior part.


represents 1/3-% of the jaw length, smaller jaws having

proportionally larger openings. In front of the opening a

narrow groove surrounds the somewhat sunk, smoothly

rounded iigament rim. The outer anteriormost margin of the rim is transforrned into a prominent, pointed, high and

narrow spur. The groove and spur are also seen in dorsal

view. A high, sharp ridge is present aiong the inner wing,

highest on its anterior part and decreasing evenly in size

posteriorly.

Left MI, dorsal side : Length 0 . 19-2 .67 mm, width slightly

less than 1/4 of length. The large , sickle-shaped fang is

slightly bent upwards in relation to the dorsal surface . The

jaw tapers continuously anteriorly from the posterior, mid

die part of the jaw. The inner margin is slightly convex,

covered by moderately large to large denticles over slightly

less than :V3 of the jaw length. The anteriormost two denti

cles are fairly small, the third usually the large st, the follow

ing 10-1 7 decreasing in size to the posterior, the dentition

ending as small knobs well before reaching the inner wing.

The dentary varies on the anterior part from paucidentate

to slightly spaced. The denticles are oriented more or less

perpendicular to extension of the jaw, though the posterior


dentary usually slants slightly towards the posterior. The

denticulated inner margin continues smoothly onto the

straight, undenticulated ridge without any notable point

transition. The inner wing is almost rectangular, fairly

small , about 1fs of the jaw length . The anterior outer end is

rounded, the inner margin parallei to the undenticulated

ridge . The basal furrow is very short, parallei to the outer

margin of the basal portion. The basal portion is fairly wide

and short, somewhat angular. The basal angle is about

30-40° . Along the anterior half of the outer margin of the

basal portion runs the ligament rim, with a conspicuous

spur. The outer margin, anterior to the basal portion, is

almost straight or with a wide and shallow concavity in its

posterior part.


represents about lf3 of the jaw length among the smaller

jaws, to 1/4 among the larger ones. The smoothly rounded,

narrow ligament rim is delimited anteriorly by a narrow

groove . Near its outer part the rim is transforrned into a

narrow, short ridge . The spur is similar to the one on the

rightjaw.

Right MIl, dorsal side : Length 0.24-2 .27 mm, width

slightly more than half the length. Two types of denticula

tion exist with only some relation to jaw size. In the first

type, two large pre-cuspidal denticles are followed by a cusp

of moderate size, only slightly larger than the pre-cusps,

and posterior to the cusp there are one or two smaller

intermediate denticles. The second type has a double cusp

with parts of almost equal size, followed by one or two

smaller intermediate denticles. The number of post-cuspi

dal denticles of both types are 8-10 , the number having no

relation to jaw size or anterior dentary. The denticles are

fairly large , slanting slightly towards the posterior, the den

tition ending before reaching the fairly pointed posterior

extremity. The shank varies from wide to fairly slender,

slightly tapering, occupying about half of the jaw length .

The inner margin is slightly convex. The deep bight has an

acute angle usually between 50° and 60° . The slender ramus

is almost triangular and has a pointed extremity. The ante

rior outer margin varies from nearly straight to slightly concave .


represents about :V3 of the jaw length . The ligament rim is

fairly narrow, the margin of the opening curves smoothly,

with an inner-anterior bulge .

Left MIl, dorsal side : Length 0 .32-l .90 mm, width

slightly less than half the length. The cusp is large and

single, or slightly smaller and double, almost equal-sized,

forming a sickle-shaped anterior margin. The intermediate

dentary is represented by 4-6 fairly large , widely spaced

denticles. The 8-1 0 fairly large, slightly slanting post-cuspi

dal denticles end at the posteriormost part of the fairly

slender shank, which occupies about half of the jaw length .

The inner wing is almost triangular, with its ,videst part

anteriorly. The bight angle varies from 60° to almost 90° .

The ramus is fairly slender and pointed, its posterior part

of ten bent to the posterior. The anterior outer margin has

a sinus.



represents about V3 of the jaw length . The myocoele open

ing is crescent-shaped, its rim fairly narrow.

Remarks. - The dentition of the MI and MIl varies from

paucidentate in some MI 's to only widely spaced denticula

tion. The right MIl has one or two pre-cuspidal denticles,

and the left MI one cusp followed by a denticle of normal

size, or a cusp followed by a large denticle . The differences

are usually consistent within and between samples and are

not regarded as be ing of taxonomic importance above the

population level . From some samples only one of the forms

has been encountered. This might be an artifact due to the

small samples. Some specimens do not fit exactly into the

description of this species and so have be en determined as

K. ( K. ) cf. burgensis (Figs. 33A-F) .

Comparison.- The basal portion of the MI is similar to the

corresponding part of K. ( K. ) polanensis cf. polanensis but

differs in having a more pronounced spur and a more

highly elevated flange . However, the size of the spur is not

a particularly good character because it is difficult to esti

mate when seen from above. The dentition is a more

conspicuous character, though in 'wide-spaced' forms the

similarity to K. ( K. ) polanensis cf. polanensis is close . The left

MIl is the best element by which to differentiate the two

species (Figs. 33A, H, 43A:3 , 44A:3, B:3 , C:3) .

Kettnerites ( K. ) huberti Bergman 1 987 Figs. l 3R, S, T, u, l SD, 34, 35 , 36

Synonymy. - 0 1 970 Paulinites polanensis Kielan-Jaworowska

1 966 - Szaniawski, pp. 465-466, Pl. I :5A-D. 0 1 987 Kettner

ites (K.) huberti n.sp. - Bergman, pp. 59-62, Figs. l 3R-U, 18 ,

34-36.

Derivation of name. - Named in honour of Dr Hubert Sza

niawski, Warsaw, who was the first to describe specimens of

the species.

Holotype. - LO 5833:7, right MI, Fig. 36H.

Type locality. - Bankvat l .

Type stratum. - Halla Beds, unit b.

Material. - Fig. 3 ; more than 250 right MI, 250 left MI, 250

right MIl, 250 left MIl.

Occurrence. - Figs . 4 and 6; Wenlock to Late Ludlow. Slite

Beds, Slite Mari , undifferentiated, to Hamra Beds, unit c .

AmIings l , Angmans 2 , Autsarve l , Bandlunde l , Bankvat

l , Barkarveård l , Bodudd l , 2, and 3, Botvide l , Djaupviks

udden 4, Faludden 2, Fie 3, Gannor l and 3, Gamudden 3

and 4, Gogs l , Grogarnshuvud l , Grundård 2, Gyle 2 ,

Hallsarve l , Hide l , Hoburgen 2 , Hummelbosholm 1 , Juves

3, Kapelludden l , .Kame 3, .Kattelviken 5, Klasård l , Kroken

2, Lambskvie l , Linviken 2, Myrsne l , Nar 2, Nisse l , Nyan

2, Ondarve l , Petsarve 2, Petsarve 15 , Ronnings l , Snau

valds l , Stora Kruse l , Strands l , Tjangdarve l , Tomt

bodame l , Traske l , Vaktård 4 and 5, Vastlaus l , Vidfalle

1 .


Diagnosis. - Right MI: Inner margin normally strongly con

vex with small denticles in fakal arch and large to very large

denticles on posterior part of the denticulated, arched

inner margin. Shank blunt-ended, flange fairly thick

walled, basal portion wide and angular.

Left MI: Inner margin convex, with somewhat smaller

denticles than right MI. Almost rectangular inner wing.

Basal portion wide and angular.

Right MIl: Two large pre-cuspidal denticles, large cusp,

large denticles on the shank, nearly parallei, slightly con

vexo-concave to almost straight sides of the shank, large,

wide, approximately triangular ramus.

Left MIl: Keel along the anterior outer margin ending in

small but distinct pre-cusp . Large, approximately triangu

lar ramus.

Description. - Right MI, dorsal side : Length 0 .32-1 .40 mm,

width V4 of length or slightly less. The inner and outer

margins are almost parallei or taper in the anterior direc

tion, ending in a fairly large fang, moderately ben t upwards

in relation to the dorsal surface . The inner margin is usu

ally convex but varies from almost straight to strongly con

vex, sometimes within one sample (e .g . the Ludlow Bank

vat l ) . The inner margin is denticulated along 0 .6--0.7 of

the jaw length, from the anteriormost part of the fakal arch

almost to the anteriormost part of the inner wing. The

denticulation is very varied. The anterior 8-1 4 denticles are

fairly small, slightly spaced, with the anteriormost denticles

directed forward. Succeeding denticles gradually increase

in size toward the central part of the dentary and then

dec re ase from the lower middle part of the posterior den

tary. This is more pronounced in the larger jaws. The

denticulation ends beside or slightly posterior to the ante

rior part of the inner wing, normally very abruptly, without

any decline in size of the posteriormost denticles (except

in, e .g . , Hallsarve l, Gannor 3 , and Bankvat 1 ) . The almost

triangular inner wing is deeply downfolded, anteriorly

rounded, widest part anteriorly, with a sometimes concave

inner margin . The undenticulated ridge is fairly large,

often bent slightly towards the outer margin, sometimes

with a smaller narrow costa parallei to and on top of the

anteriormost part of the ridge . The ridge, the extremity of

which is bent towards the inner margin, forms the posterior part of the jaw, the blunt-ended shank. The basal furrow is of normal length and width . The flange is fairly thickwalled, very varied in shape from angular with the outer margin bent down (Fig. 36J, Bankvat l , Hamra Beds) to

rounded (Fig. 34E, Gannor 3, Hemse Beds) , with a furrow

along its outer margin of the basal portion and its outer

most margin folded upward. Much, but not all , of the

variation depends on the preservation, i .e . compaction of

the jaws increases the size of the more or less pronounced

folds and furrows. The basal portion is wide and distinct,

but there is a large variation in shape. The basal angle is low

to very low, almost zero. The outer margin is normally

concave, but variation occurs from straight to strongly

concave, especially in the Ludlovian material . The outer

margin ends in a rounded falx.

Ventrai side: The strongly enclosed myocoele opening

occupies about V3 of the jaw length. The anterior margin is



Fig. 35. Kettnerites (K) huberti. All specimens in dorsal view, x 1 20, except B4. DA. Vaktård 4, Hemse Beds, Hemse Mari, SE part, 81-35LJ; Al left MI, LO 5822 : 10 ; A2 right MI, LO 5822: 1 1 ; A3 left MIl, stereopair, LO 5822: 1 2; A4 right MIl , stereopair, LO 5822: 13 . DB. Strands l , Hamra Beds, unit b , 75-14LJ; B 1 left MI, LO 5840: 1 ; B 2 right MI, LO 5840:2 ; B3 left MI, LO 5840:3; B 4 right MI, LO 5840:4, x80.

crescent-shaped and surrounded by the ligament rim, a roll-shaped ridge which in turn is separated from the ante

rior of the jaw by a narrow groove . In the anteriormost part

of the inn er wing the ridge forms a narrow, high (in lateral

view, triangular) proeess. Left MI, dorsal side: Length 0.30-l .52 mm, width less

than 1/4 of length . The jaw tapers anteriorly, ending in a

Fig. 34. Kettnerites (K) huberti. All speeimens in dorsal view, x1 20. DA. Kårne 3, Eke Beds, lowermost part, 71-198LJ; Al left MI, LO 5843 :4; A2 right MI, LO 5843:5 . D B. Gannor l, Eke Beds, lowermost part, 71-123LJ; Bl left MI, LO 581 1 :5; B2 right MI, LO 581 1 :6. D e. Hide l, Slite Beds, Slite Mari, 73-2LJ; Cl left MI,

LO 5784: 1 ; C2 right MI, LO 5784: 2; C3 left MIl, LO 5784:3; C4 right MIl, LO 5784:4. OD. Hallsarve l, Hemse Beds, Hemse Mari, uppermost part, 69-28LJ; Dl left MI, LO 5803 : 1 ; D2 right MI, LO 5803:2 . DE. Gannor 3, Hemse Beds, Hemse Mari, SE part, 71-1 25LJ; E1 1eft MI, LO 58 14: 1 ; E2 right MI, LO 5814:2 .

fairly large fang, bent moderately upwards relative to dorsal surface . In a few jaws, the inner and outer margins are

almost straight and parallei, though the inner margin is

normally distinetly convex. Denticulation covers 0 .5-0 .7 of

the jaw length, from the anteriormost part of the falcal arch to near the anteriormost part of the inner wing. The denti

des in the falcal arch range in number and size from about

5 moderately large, widely spaced dentides to about 15

small ones, the latter type being more common. In some

jaws the dentides are more or less widely spaced crenula

tions on a low ridge . Posterior to the falcal arch follow

larger slanting dentides, though the difference in size

between the anterior and posterior on es is less than in the

right MI. The denticulation ends abruptly or with some

small dentides. The undenticulated ridge is low and fairly

narrow, forming a smooth, straight continuation of the

denticulated inner margin. The somewhat downfolded



inner wing is fairly large, 0 .25-0.30 of the jaw length, alm ost

rectangular with its inner margin straight to slightly con

vex, and oriented parallei to the denticulated ridge or with

the anterior corner pointing away from the latter. The

basal furrow is fairly long and narrow. The posterior mar

gin is long and almost straight. The outer margin of the

basal portion represents almost 0 .3 of the jaw length and is

convex to straight with a small er 'outer wing' on its anter

iormost part. The basal portion is wide, angular and prom

inent. The basal angle varies but is usually low. The outer

margin in front of the basal portion is more or less concave,

ending in a large, sickle-shaped falx.

Ventrai side: The strongly enclosed myocoele opening,

occupies about V3 of the jaw length . The anterior margin

of the opening is crescent-shaped, drawn slightly forwards

along the inner margin of the jaw. A fairly narrow, rounded

ridge is surrounded by a groove . The ridge forms a small

process on the anterior part of the inner wing and a sharp

edge along the ventrai inner margin of the inner wing.

Along the outer margin it is unfolded.


about half the length or slightly less. Two large pre-cuspidal

denticles, with distinct cutting edges, form a large protrud

ing anterolateral part of the jaw, with an acute to right-an

gled margin. The cusp is fairly large and posteriorly di

rected; it is followed by 2-3 intermediate denticles

increasing in size towards the posterior, and normally 6--8

fairly large succeeding denticles. All denticles along the

slightly convex to almost straight inner margin slant. The

inner wing is very narrow. The shank is long, normally

slightly more than half the jaw length, with convexo-con

cave to almost parallei sides. The posterior extremity of the

Wenlock specimens is directed outwards (Fig. 34C:4) ,

while the termination on the Ludlow specimens is more or

less straight and blunt. The shank is also slender among the

Wenlock specimens. The bight angle varies among differ

ent populations, from acute (Fig. 34C:4) to almost right

angled (Figs. 35A:4, 36D) . The ramus is fairly short and

broad, the anterior outer margin has a more or less pro

nounced sinus.


represents �3-% of the jaw length . The ligament rim is

narrow, slightly curved, the opening extending forwards along the inner margin .

Left MIl , dorsal side: Length 0.24-2 .03 mm, width about

half the length . A small but distinct pre-cuspidal denticle

forms the anteriormost part of the jaw; its continuation is

represented by a characteristic smooth keel running along

the anterior outer margin and disappearing in the sinus at

Fig. 36. Kettnerites (K.l huberti. Bankvat l , Hamra Beds, unit b. C-G from sample 82-34CB, the remaining ones from 81-39LJ. All speei

mens are in dorsal view, A-G xl20 and H-O x80. O A. Left MIl, LO 5833:9. DB. Right MIl, LO 5833: 10 . OC. Left MIl, LO 5834:3 . OD. Right MIl , LO 5834:4. DE. Left MI, LO 5834:5. OF. Left MI, LO 5834: 1 . OG. Right MI, LO 5834:2. OH. Holotype, right MI, LO 5833: 7. DI . Right MI, LO 5833:8 . OJ. Right MI, LO 5833:2. D K. Left MI, LO 5833:3. D L. Left MI, LO 5833:6. DM. Left Ml, LO 5833: 1 . ON. Left MI, LO 5833:5. DO. Left MI , LO 5833:4.


the base of the ramus. The cusp is of normal size, usually

followed by 4-6, sometimes 8-1 0, intermediate denticles of

equal size. The size of these denticles varies among the

different populations. The inner wing is fairly short and

rounded. The inner and outer margins of the shank are

almost parallei and the shank represents somewhat less

than half of the jaw length . Normally the posteriormost

part of the shank projects posteriorly, though in the Wen

lock material, a slightly outward direction is normal. The

bight angle is more or less 90'. The ramus is fairly short with

a distinct sinus in the outer margin at the basal part.


represents about half of the jaw length . The slightly cres

cent-shaped opening is surrounded by a narrow ligament

rim which gets wider at the outer side .

Remarks. - Wenlock and Lower Ludlow populations seem

to be less varied than those of the Middle and U pper

Ludlow. This could be an effect of the very limited material

from the older strata. The population from the Middle

Wenlock locality, Hide l, is very similar to the normal

shaped specimens from, e .g . , the Upper Ludlow locality

Bankvat 1 . However, the high number of intermediate and

post-intermediate denticles found among some left MIl

from the Ludlow, distinguish them from the normal form.

I suspect that the left MIl with the larger num ber of denti

cles (Fig. 36C) should be grouped with the slender MI

(Figs. 35A: I , B:3) . Thus, the species seems to show a phylo

genetic trend towards at least three different morphs. The

normal morph has an MI with a pronounced convex inner

margin carrying large, spaced denticles (e .g. Figs . 34C: I ,

C :2 , E : I , E :2 , 36H, I , K, M) . The second morph has a less

wide and almost straight MI, and slightly smaller denticles

(Figs. 36F, G, J) . The third, less common, morph has a

slender s-shaped left MI and a right MI with reverse s-shape

and smaller denticles. As late as in the Late Ludlow, the

morphs can be found in the same samples. This implies

that they are still to be regarded as one species.

Discussion. - This species seems to have been widely spread

in the shallow sea of Baltica. The Upper Ludlow species

from the Mielnik bore hole in Poland (identified by Sza

niawski 1 970 as Paulinites polonensis Kielan:Jaworowska) , known from three apparatuses and isolated elements, is very similar to the Gotland species and is probably conspe

cific with the latter. The type specimen of Oenonites aspersus

(Hinde 1 880, p. 373, Pl. 1 4:7) is compressed and partly

buried in the rock; only a very slight resemblance to K. ( K. ) huberti may b e noted. Kettnerites (K.) huberti i s also present

in the Ludlow of the Welsh borderland (courtesy Alison

Brooks) .

Comparison. - Both the left and right MI of the normal

morph (i .e . having a convex inner margin with large denti

cles) are characteristic and easy to identifY. The second

morph is less heavily denticulated and resembles the corre

sponding elements of K. (A . ) sisyphi klasaardensis, but dif

fers in having small denticles in the anterior part and its

largest denticles in the middle part of the denticulated

inner margin. The basal portion is also slightly different;

70 Claes F. Bergman

usually the basal plate is laeking on the right MI of K. ( K. ) huberti.

The right MIl has two pre-cuspidal dentides, which is

fairly common within Kettnerites, e.g . , K. ( K. ) polonensis, K. ( K. ) bankvaetensis, and some of K. (K.) burgensis. The pre

cuspidal dentides of K. (K.) huberti are of equal size, in

contrast to those of K. ( K. ) bankvaetensis. Further, the ram us

of the MIl of K. (K.) huberti is large and almost triangular,

which distinguishes the speeies from K. ( K. ) polonensis and

K. (K.) burgensis, these having a more or less needle-shaped

ramus (Figs. l 8A, B, D, E) .

Kettnerites ( K. ) martinssonii Bergman 1987 Figs. l 2A-G, l 8C, 37-40

Synonymy. - O 1 882 Eunicites cristatus (Hinde 1 879) - Hinde,

p. 10, Pl. 1 :6, right MIl. 0 1 882 Arabellites anglicus Hinde

1 880 - Hinde, p. 18 (pars. ) , Pl. 2 :50, left MIl. 0 1 882

Oenonites aspersus Hinde 1 880 - Hinde, p. 13, Pl. 1 : 2 1 , 22,

left MI. 0 1 960 Paulinites sp. - Martinsson, pp. 5-6, Fig. 1 :6,

a jaw apparatus. 0 1 979 Oenonites aspersus Hinde - Berg

man, p. 99 (pars. ) , Fig. 28:4B, E, F. 0 1987 Kettnerites ( K. ) martinssonii n.sp. - Bergman, pp. 62-66, Figs. 1 2A-G, 1 8,

37-40.

Derivation of name. - In honour of the late Professor Anders

Martinsson, who increased our knowledge of the geology

and biostratigraphy on Gotland considerably and was the

initiator and coordinator of the Project Ecostratigraphy.

He was also the first to describe polychaete apparatuses

from Gotland. He found two apparatuses and named one

but left this species under open nomendature . This appa

ratus belongs to the very common speeies now named

Kettnerites (K.) martinssonii.

Holotype. - LO 5764: 1 , right MIl, Fig. 37G.

Type locality. - Vattenfallsprofilen l .

Type stratum. - Hogklint Beds, unit b.

Varieties. - K. ( K. ) martinssonii var . mulde .

Material. - Figs. 2 and 3; more than 250 elements each of right MI, left MI, right MIl, left MIl.

Occurrence. - Figs. 4 and 9; Latest Llandovery to Late Ludlow, Lower Visby Beds to Hamra Beds. Ajmunde l , AmIings

1 , Ansarve l , Ar l , Aursviken 1 , Autsarve l , Baju 1 , Band

lunde 1 , Blåhall 1 , Bodudd l and 3, Bofride l , Bottarve 2 ,

Buske l , Dapps l and 2 , Djupvik 1 , 2 , 3 , and 4, Fardume 1 ,

Fie 3 , Fjale 3 , Fjardinge 1 , Follingbo 2 and 1 2 , Gamla Hamn

1, Gandarve 1 and 2 , Gannor 1 and 3, Garnudden 3 and 4,

Gerete l, Gerumskanalen l, Glasskar 1 and 3, Glaves 1 ,

Gnisvard 1 and 2 , Gogs l , Gothemshammar 1 , 2 , 6 and 7,

Grogarnshuvud l, Grymlings 1 , Gustavsvik 2 , Gutenviks 1 ,

Gyle l and 2 , Hiiftingsklint l and 4 , Haganas l , Hagur l ,

Hallagrund 1 , Halls Huk l , Hide 1 , Hide Fiskelage l ,

Hoburgen 2 , Horsne 3 and 5 , Hummelbosholm l , Ireviken

2 and 3, Kappelshamn 1 , Kattelviken 5, Kauparve l , Klinte

berget l , Kluvstajn 2, Korpklint 1 , Kroken 2, Kullands l and

2, Langhammarsviken 2, Lauter 1 , Lauterhornsvik 2, Lick

ershamn 2, Lilla Hallvards l and 4, Loggarve 2, Mollbos l ,


Molner l , Mulde 2, Mulde Tegelbruk 1 , Nabban 2 , Nar 2 ,

Narshamn 2 and 3 , Nygårds l and 2 , Nygårdsbii.ckprofilen

l , Nyhamn 4, Ondarve 1 , Paviken 1 , Petsarve 2 and 1 5 ,

Rågåkre l , Rangsarve l , Ronnings l , Ronnklint l , Saxriv l ,

Sigvalde 2 , Slitebrottet l and 2 , Snackgardsbaden l , Snoder 2 and 3, Sproge 4, Stave 1 , Stora Myre 1 , Storugns l B,

Strandakersviken l , Stutsviken 1 , Svarvare l , Svarven l ,

Talings l , Tjeldersholm 1 , Tomtbodarne 1 , Traske 1 , Val by

bodar 1 , Valle 2, Valleviken 1 , Vallmyr l , Vallstena 2 , Valve

3, Varsande l , Vastlaus l , Vattenfallsprofilen l , Vidfalle 1 ,

Vike 2 and 3.

Diagnosis. - Right MI: Subtriangular, sharp ly ended shank,

thin-walled, narrow, highly elevated flange . Almost triangu

lar, small basal portion .

Left MI: Dentides of normal size to fairly large, slightly

to widely spaced in anterior half, somewhat posteriorly

directed, almost rectangular inner wing.

Right MIl: One large pre-cuspidal dentide, cusp of mod

erate size. Ramus fairly large , almost triangular, wide in

basal part, anterior outer margin nearly straight. Shank

fairly wide, tapers to posterior.

Left MIl: Single cusp, followed by 5-8 intermediate den

tides of equal size. Ramus fairly large, almost triangular,

wide in basal part.

Fig. 37 (opposite page) . Kettnerites (K.) martinssonii. All speeimens in dorsal view, x1 20, except B2, E and 1 1 . DA. Buske l, Lower Visby Beds unit e, 79-40LJ; Al left MI, LO 5362: 1 ; A2 right MI, LO 5362 :2 . DB. Follingbo 2, Slite Beds, Slite Mari NW part, 75-1 0CB; B l left MI, LO 5783: 1 ; B2 right MI, LO 5783 :2, x80. OC. Right MIl, LO 5767: 1 , Vattenfallsprofilen l , Hogklint Beds, unit b, 70-6L]. OD. Left MIT , LO 5763: 1 , Ansarve l , Hogklint Beds, SW facies, upper part, 79-46L]. DE. Right MIl, LO 5767:2, Vattenfallsprofilen l , Hogklint Beds, unit b, 70-6LJ, x80. OF. Left MIl, LO 5771 : 1 , Valle 2, Slite Beds, Pentamerus gothlandicus Beds, 6f>l45SL. O G. Holotype, right MIl, LO 5764: 1 , Vattenfallsprofilen l , Hogklint Beds, unit a , 70-20L]. O H . Vattenfallsprofilen l , Hogklint Beds, unit b, 70-6LJ; Hl left MI, LO 5767:3 ; H2 right MI, LO 5767:4. O I . Vattenfallsprofilen l , Hogklint Beds, unit a, 70-20LJ; 11 left MI, LO 5764:3, x80; 12 right MI, LO 5764:2, x80. O]. Slitebrottet 2 , Slite Beds Slite Mari, 83-3lLJ; J l left MI, LO 5773: 1 ; J 2 right MI, L O 5773 :2 .

Fig. 38 (p . 72) . Kettnerites (K.) martinssoniivar. mulde. All specimens in dorsal view. DA. Mulde Tegelbruk l, Mulde Beds, 82-7CB; Al right MIl, LO 5789: 1 , x120 ; A2 left MIl, LO 5789:2 , x1 20. D B.

Dapps l , Mulde Beds, upper part, 81-56LJ; B l left MI, LO 5787: 1 , x80; B 2 left MI, LO 5787:2, x80; B 3 right MI, LO 5787:3, x80; B4 right MI 5787:4, x80 ; B5 left MIl, LO 5787:5, x60; B6 right MIl,

LO 5787:6, x60.

Fig. 39 (p . 73) . Kettnerites (K.) martinssonii. D and E are K. (K.) martinssonii var. mulde . A , C, E: Snoder 2 , Hemse Beds, Hemse Mari, NW part, 82-14CB. B, D : Kullands 2, Hemse Beds, Hemse Mari, NW part, 84-31 2DF. All speeimens in dorsal view. All x 120, except E. DAl , left MIl, LO 5808:8; A2 right MIl, LO 5808:7 . OBl right MI, LO 5832: 2 ; B2 left MI, LO 5832: 1 . OCI left MI, LO 5808:3; C2 right MI, LO 5808:4. OD! left MI, LO 5832:3; D2 right MI, LO 5832:4. DEl left MI, LO 5808: 1 , x80; E2 right MI, 5808:2, x80; E3 left MIl, LO 5808:5, x80; E4 right MIl, LO 5808:6, x60.

FOSSILS AND STRATA 25 ( 1 989) Silunan paulinitid polychaetes from Gotland 7 1







Description. - Right MI, dorsal side: Length 0 .43-l .23 mm

(mulde variety 0 .65-2 .28 mm) , width lf3-lf4 of length. The

jaw tapers an teriorly, ending in a normal to fairly large fang

bearing a fairly distinct cutting edge . The inner margin is

convex, denticulated with about 30 denticles of almost

equal size in Early Wenlock specimens. During the Middle

Wenlock to Late Ludlow, the denticles increase in size,

particularly the anterior ones, and the num ber decreases

to about 20. The dentition covers the inner margin from

the falcal arch, ending, with slightly spaced denticles, be

fore reaching the inn er wing. The large jaws from the Late

Wenlock strata of ten have large to very large, slightly

spaced denticles. The continuation of the inner margin

onto the undenticulated ridge is smooth . The undentic

ulated ridge is long, high and narrow, its middle part

slightly bent outward, ending as a prominent tip. The tip

forms the sharp-ended shank together with the posterior

part of the inner wing. The inner wing is of normal size,

deeply downbent, wide , its anterior half rounded. The

basal furrow is deep and long. The flange is thin-walled,

high ly elevated, narrow, ending postero-Iaterally as a

slightly downbent tip or blunt end. The flange passes over

anteriorly into the fairly large and rounded ligament rim.

This rim has a rough surface and forms a more or less

pronounced ridge , sometimes with a gro ove on its inner

side. A conspicuous, wide, outer border is present in the

mulde variety from the late Middle Wenlock ( see Re

marks) .


The basal portion is wide and narrow, alm ost triangular,

with a basal angle of about 35-50' . The posterior part of the

jaw is widest, with a concavity in its middle part whence it

extends forming the sickle-shaped falx.

Ventrai side : The strongly enclosed, crescent-shaped

myocoele opening represents one third of the jaw length.

It is surrounded by a fairly narrow ligament rim.

Left MI, dorsal side: Length 0 .40-l .54 mm (mulde vari

ety 0 .73-2 . 1 8 mm) , width lf3-1f4 of length. The inner mar

gin is almost straight (Figs. 37H: 1 , 39B:2) to convex with

fairly large, slightly spaced denticles on the anterior part

(Figs. 39D : 1 , E : 1 , 40A: 1 , B) . The anteriormost denticles

may be oriented anteriorly. Denticles, about 30 in num ber

in Early Wenlock specimens, decrease in number but in

crease in size with stratigraphic time . In Ludlow specimens the anterior denticles are large to very large and the poste

rior ones are small knobs. Their number varies greatly but

Fig. 40. Kettnerites (K) martinssonii. D are K (K) martinssonii var. mulde. All speeimens in dorsal view. DA. Glasskar 3, Burgsvik Beds, 82-18CB; Al left MI, LO 5828: 1 , x80; A2 right MI, LO 5828:2, x80. DB. Left MI, LO 5844:3, Kauparve l, Hamra Beds, lower-middle part, 76-1 3CB, x80. DC. Gannor l, Eke Beds, 7l-1 23LJ; Cl left MI, LO 581 1 : 1 , x1 20; C2 right MI, LO 58 1 1 :2 , x1 20. OD. Varsande l , Mulde Beds, 75-22CB; D l left MI, LO 5793: 1 , x1 20; D 2 right MI, LO 5793:2, x80; D3 left MIl, LO 5793:3, x80; D4 right MIl, LO 5793:4, x80. DE. Sigvalde 2, Hemse Beds, lower middle part, 71-1 15LJ; El left MI, LO 5819: 1 , x 1 20; E2 right MI, LO 5819:2 , x1 20. O F. Traske l , Hemse Beds, unit b, 7 l-1 03LJ; Fl left MI, LO 581 8 : 1 , x60; F2 right MI, LO 5818:2, x60; F3 left MIl , LO 5818:3 , x120; F4 right MIl, LO 581 8:4, x60.


is normally about 20. The denticulation ends as crenula

tions well before reaching the inner wing. The latter is

slightly downfolded, of ten posteriorly projecting, almost

rectangular, nearly angular, but highly variable in shape.

The denticulated inner margin has a smooth and straight

continuation onto the low and narrow undenticulated

ridge , which ends before reaching the posterior margin,

with a low ridge oriented towards the posterior, inner

corner of the inner wing. The basal furrow is of normal size.

The posterior margin is short and concave . The posterior

part of the outer margin of the basal portion is downfolded

and has a fairly narrow flange on its anterior part. The basal

portion is fairly small, the basal angle usually 30-40' . The

outer margin between the basal portion and the large flat

to rounded falx is concave . The cutting edge of the fang is

fairly distinct.


myocoele opening represents 1/4-1!� of the jaw length . The

opening is very similar to the one on the right MI.

Right MIl, dorsal side : Length 0.35-l . 1 3 mm (mulde

variety 0 .38-l .80 mm) , width more than half the length . A

large , pre-cuspidal denticle with a distinct cutting-edge,

forms the anteriormost, fairly acute margin of the jaw. It is

followed by a somewhat larger, moderate-sized cusp, di

rected slightly to the posterior or almost perpendicular to

the denticulated ridge . Posterior to the cusp there are one

or two minor denticles of which the anteriormost is the

smaller, followed by 6-9 large to very large denticles, slant

ing towards the posterior direction and decreasing slightly

in size towards the posterior; the posteriormost ones are in

some cases very small. The number of denticles is indepen

dent of jaw length. The anterior part of the shank is wide ,

tapering slightly towards the posterior, and occupies

slightly more than half of the jaw length . The ramus is fairly

large, usually without any sinus in the outer margin, almost

triangular, wide in the basal part. The bight angle is 55-75' .

Ventrai side: The slightly enclosed, crescent-shaped myo

coele opening is lf2-:V3 of the jaw length . The opening is

extended slightly forwards along the inner margin. The

ligament rim is narrow.

Left MIl, dorsal side: Length 0.33-l .22 mm (mulde vari

et y 0 .38-1 .73 mm) , width about half the length or slightly

less. The single cusp with its distinct cutting edge forms the anteriormost part of the jaw. The intermediate dentary has 5-8 denticles of equal and moderate size , followed by 8-1 0

larger denticles, reaching maximum size i n the central part of the dentary. The number of denticles is unrelated to the

size of the jaw. The inner wing is pronounced, representing about half of the jaw length . The inner margin is almost

parallel with the denticulated ridge . The shank occupies

about half or slightly less of the jaw length, its anterior part

being very wide and tapering towards the posterior, ending

in a fairly sharp tip . The ram us is fairly large, almost trian

gular, wide in the basal part, with a sinus in the anterior

outer part. The bight angle is 60-85' .


represents slightly more than half the jaw length . The

anterior margin is somewhat curved with its main orienta-

76 Claes F. Bergman

tion oblique to the extension of the jaw. The ligament rim

is narrow.

Remarks. - During the latest Llandovery and Early to Middle

Wenlock the MI jaws of this speeies are usually 0 .5-1 mm in

size. The shape of the jaws varies slightly from population

to population. The inner margin is usually almost straight

in the left MI. During the later part of the Wenlock and in

the Early Ludlow, i .e . Halla, Mulde , Klinteberg, and Hemse

Mari , NW part, however, the maximum size of the jaws

increased and a new shape of the larger MI evolved (Fig.

1 2A-G) . The inner margin became pronouncedly convex,

the flange grew larger, the outer anterior part of the flange

became conspicuously wider, forming a border (e .g . Fig.

38B:1-4) . I have named this variety 'mulde ' after the Mulde

Beds where these characteristics become most conspicu

ous.

During the Middle and Later Ludlovian the maximum

size seems to have decreased slightly and the jaws became

more curved and somewhat thinner, with comparably

larger denticles. The posterior, outer margin of the basal

portion of the left MI is normally considerably shorter and

slightly concave (Fig. 40A, B, and C) .

The dentition formula and the morphology of the MIl

are virtually unchanged from the latest Llandovery to Late

Ludlow. The size changes follow those of the MI.

Comparison. - The right MIl with its large pre-cuspidal

denticle , followed by a fairly large cusp is the best character

for identification. Large MI ' s younger than the midd le

Wenlock, with their large dentary, convex inner margins

and conspicuous basal portion, are also easy to identify.

The Left MIl, however, is considerably more difficult to

identify. The corresponding element of some of the forms

of Kettnerites (A. ) sisyphi is virtually impossible to distinguish

from K. ( K. ) martinssonii. There is no safe character to rely

on, though the cusp of the left MIl of K. (A. ) sisyphi is

normally slightly wider and somewhat flattened obliquely

to the extension of the jaw. The K. ( K. ) martinssonii left MIl

jaw might give a slightly coarser impression and is often

larger than in K. (A . ) sisyphi. Comparisons with K. ( K. ) bankvaetensis were given above .

Kettnerites kosoviensis Zebera from the Silurian of Bohe

mia is probably closely related to K. ( K. ) martinssonii. They are probably not conspecific. Zebera's speeies ( 1 935) was

revised by Snajder ( 1951 ) who collected new material from the same and other localities used by Zebera, and described

the polychaete fauna there according to a biological spe

eies concept. The badly preserved elements from the

Bohemian differ from the Gotland ones in that the number

of denticles of the MIl is somewhat higher and the denticles

of the MI seem to be thinner. However, the jaws from

Bohemia are very large, flattened and crushed, making a

comparison very difficult. New material from Bohemia,

showing three-dimensional morphology and variability, is

needed to resolve this question.


Kettnerites (K) polonensis (Kielan:Jaworowska 1 966) Figs. 1 3A-C, 1 8A, 4 1 , 42B, E, 43 , 44

Synonymy. - 0 1 980 Paulinites polonensis Kielan:J aworowska

1 966 - Wolf, pp. 86--87, Pl. 1 2 : 1 02 . 0 1 987 Kettnerites ( K. ) polonensis (Kielan:Jaworowska 1 966) - Bergman, pp . 66--69,

Figs. 1 3A-G, 1 8, 4 1 , 42B, E, 43, 44.

Holotype. - 'Paulinites ' polonensis Kielan:Jaworowska 1 966,

pp. 1 26--1 29, Pl. 29 :2 .

Type locality. - Erratic boulder.

Type stratum. - Not known, see discussion below.

Varieties. - K. (K. ) polonensis var. gandarve, K. ( K. ) polonensis

var. sjaustre .

Material. - Fig. 3; more than 250 right MI, 250 left MI, 250

right MIl, 250 left MIl .

Occurrence. - Figs. 4 and 1 0; Early Wenlockian to Late

Ludlovian, Hogklint Beds to Sundre Beds. AmIings l , Baju

l, Bankvat l, Barkarveård l, Botvide l, Djaupviksudden 4,

Fågelhammar 3 , Fakle l, Faludden 2 , Fjardinge l, Gannes

2 , Gannor l and 3, Garnudden 1 , 3 , and 4, Gerumskanalen

l , Glasskar l , 2, and 3, Glaves l , Godrings l and 2, Gothems

hammar l , 2, 6, 7 and 8, Grogarnshuvud l , Grondalen l , Gyle l and 2 , Hallsarve l , Herrvik 2 , Hide l , Hoburgen 2 ,

Holmhallar l , Kapelludden l , Kattelviken 5 , Klinteberget

l , Kroken 2, Kullands 2, Lambskvie l , Loggarve l and 2 ,

Mollbos l and 2 , Mulde 2 , Mulde tegelbruk l , Nar 2 ,

Narshamn 2 , Nyan 2 , Nygårds 2 , Rågåkre l , Ronehamn 2 ,

Rudvier l , Sibbjans 2 , Sjaustrehammar l , Slitebrottet 2 ,

Sigvalde 2 , Stora Vikare 2 , Strandakersviken l , Strands l , Tradgården l , Traske l , Valby Bodar l , and Vasterbackar

1 .

Fig. 4l. Kettnerites polonensis Kielanjaworowska 1 966, camera-lucida drawing of the type specimen apparatus consisting of the right MI, left and right MIl (Z. Pal . No. 0.439/3) , x235. The MIII element was not fused to the apparatus in 1 985. Note the following charac

teristics: the right and left MI have almost rectangular inner wings and prominent spurs; the right MIl lacks the pronounced cusp and left MIl has a double cusp.


Fig. 42. Kettnentes (K.) polonensis var. gandarve, except B and E which are K. (K.) polonensis. All specimens are in dorsal view. DA. Sjaustrehammar l, Hemse Beds, unit d, 82-l9LJ; Al left MI, LO 58 1 6: 1 , x 170; A2 right MI, LO 581 6:2, x 1 70; A3 left MIl, LO 58 1 6:3 , x 120 ; A4 right MIl, LO 581 6:4, x1 20. DB . Right MI , LO 581 3:3, Gannor 3, Hemse Beds, Hemse Mari SE part, 71-126LJ, x1 20. DC. Gandarve l , Halla Beds, 7 1-81 LJ; Cl left MI, LO 5836:4, xl OO; C 2 right MI, L O 5836:5, xl OO; C3 left MIl, LO 5839:2 , x1 20; C4 right MIl, LO 5839:3 , xl00 . OD . Gothemshammar 7, Halla Beds, unit c , 77-45LJ; Dl left MI , LO 5837: 1 , x80; D2 right MI , LO 5837:2, x80; D3 left MIl, LO 5837:3, x80;

D4 right MIl, LO 5837:4, x80. DE. Gannor l, Eke Beds lower part, 71-l24LJ; El left MI, LO 581 2 : 1 , x80; E2 right MI, LO 581 2:2, x80.


Fig. 43. Kettnerites (K.) polonmsis. All specimens are in dorsal view. DA. Glasskar 3, Burgsvik Beds, lowermost part, 82-18CB; Al left MI, LO 5827:3, x60; A2 right MI, LO 5827:4, x60; A3 left MIl, LO 5827:5, x60; A4 right MIl, LO 5827:6, x60. DB. Lambskvie l, Hemse Beds, unit c,

75-45CB; Bl left MIl, LO 5817: 1 , x60; B2 right MIl, LO 5817:2 , x60. O C. Left MI, LO 5842: 1 , Glasskiir l, Burgsvik Beds, lowermost part, 82-1 5CB, x80. OD. Same sample as C; Dl left MI, LO 5842 :2, x80; D2 right MI, x60.

Fig. 44 (opposite page ) . Kettnerites (K.) polonmsis. All specimens in dorsal view. DA. Faludden 2 , Hamra Beds, unit c, 76-16CB; Al left MI, LO 5830: 1 , x120; A2 right MI, LO 5830:2, x1 20; A3 left MIl, LO 5830:3, x210; A4 right MIl, LO 5830:4, x210 . DB. Sibbjans 2, Hamra Beds, unit b, 82-32LJ; Bl left MI, LO 5845: l, x120; B2 right MI, LO 5845:2 , x 120 ; B3 left MIl, LO 5845 :3, x 1 70, B4 right MIl, LO 5845:4, x1 70. OC. Vasterbackar l , Sundre Beds, middle-upper, 75-2LJ; C l left MI, LO 5838: 1 , x80 ; C2 right MI , LO 5838 : 1 , x60; C3 left MIl , LO 5838:2 , x80; C4 right MIl, LO 5838:3, x210 . OD. Left MI, LO 5845 :5, Sibbjans 2, Hamra Beds, unit b , 82-32LJ, x60.


80 Claes F. Bergman

K. (K.) polonensis var. gandarve (Fig. 42A, C, D) is re

corded from the following Middle to Late Wenlock Iocali

ties (not included in the K. (K.) polonensis list) : Fardume l , Gandarve l , and Gothemshammar 7.

Diagnosis (emended) . - Right MI: Large fang, inner margin

slightly convex, covered by denticles of varying size. Basal

portion angular, shank ending bluntly, flange angular and

thick-walled with a gro ove and a spur on the anterior outer

part.

Left MI: Convex inner margin with large denticles ante

riorly, decreasing in size posteriorly. Basal portion angularly square with a groove along its outer margin . Prominent rectangular inner wing.

Right Mil: Two large pre-cuspidal denticles of equal size; shank long, straight and blunt-ended. Ramus needle-Iike,

long, straight, narrow, with pointed extremity.

Left Mil: Large double cusp of equal size; shank large and almost straight. Ramus needle-like, long, straight, nar

row, with pointed extremity.


width about 1/4 of length . The jaw tapers, ending in a large

fang, bent upwards in relation to jaw surface; the lower

middle part of the inner margin is slightly convex. The

denticulation covers 0 .60-0.65 of the jaw length . The large

anteriormost denticle is followed by about five denticles

that decrease to fairly small size . Posterior to these are 5-8

denticles which slowly increase in size, followed by a series

of about 10 large denticles. The posteriormost denticles are

of ten small, reduced to indistinct knobs, probably due to

wear. Rarely, jaws with denticles of more or less even size

occur. The undenticulated ridge is large and slightly sig

moidal, ending in the posteriormost part of the bluntly


Fig. 45. Kettnerites (K) cf. polonensis. All speeimens are in dorsal view. DA. Left MI, LO 5827 : 1 , Glasskar 3, Burgsvik Beds, 82-1 8CB, x80. DB. Right MI, LO 5827:2 , same sample as A, x80. D e. Right MI, LO 5842 :3, Glasskar l , Burgsvik Beds, 82-15CB, x80.

ended shank. The deeply downfolded inner wing is of

normal size, sigmoidal, and alm o st paralleI to the undentic

ulated ridge . The basal furrow is fairly narrow; the thick

walled, rectangular flange is highly elevated along the pos

terior margin and downbent along the outer margin . The

ligament rim on the posterior outer margin of the basal

portion is about 0 . 1 5 of the jaw length and consists of a

longitudinal, narrow groove with a small spur on its anter

iormost part.

The basal plate is rarely recorded in place, except some

times in the gandarve variety (Fig. 42A:2 ) and in K. ( K. ) cf.

polonensis (Fig. 45B) . It is somewhat triangular, longer than high, and with a high, convex inner margin .

The basal portion is almost triangular and the basal angle

usually 1 3-30° . The outer margin, anterior to the basal portion , is almost straight, ending in the prominent sickle

shaped falx.

VentraI side : The strongly enclosed crescent-shaped myocoele opening is about lf3 of the jaw length . In front of

the opening a narrow gro ove surrounds the somewhat

sunk, smoothly rounded ligament rim. The outer, anterior

most margin of the rim forms a prominent, short, pointed,

high and narrow spur. The gro ove and spur are visible also

in the dorsal view.

Left MI, dorsal view: Length 0.43-2 .60 mm, width about

lf3 of length . The large fang is relatively strongly bent

upwards in relation to the jaw surface . The jaw tapers

anteriorly from the posterior middle part. The inner mar

gin is slightly convex, covered by moderately large denticles

for about 0 .5-0.6 of the jaw length. Normally the size of the

20-25 denticles decreases posteriorly in a regular way. At

some localities (e .g . Glasskar l , Sibbjans 2) the anterior

most 1-2 denticles are large and followed by some 1 0 small


denticles, which in turn are followed by 8-1 2 large ones.

The denticulated inner margin continues smoothly onto

the straight undenticulated ridge which does not reach the

posterior margin. The inner wing is rectangular, pro

nounced, fairly large, 0 .25-0.37 of the jaw length, paralleI

with the undenticulated ridge . The basal furrow is short

and fairly wide, paralleI to the outer margin of the angular,

nearly rhombic, basal portion. The basal angle is medially

about 20° (varies between 15 ° and 30° ) . The ligament rim

runs along the anterior half of the outer margin of the basal

portion, forming a spur, followed laterally by a groove . The

outer margin anterior to the basal portion has a very vari

able outline (cf. 42C : l and 43A: l ) .

VentraI side: The strongly enclosed, crescent-shaped

myocoele opening represents about 1J4-1J3 of the jaw

length. The smoothly rounded narrow ligament rim is

surrounded anteriorly by a narrow groove . Toward its outer

part the rim is transforrned into a narrow ridge similar to

the one on the right MI.

Right MIl, dorsal side: Length 0.38-2 .05 mm, width

somewhat more than half the length. Two pre-cuspidal

denticles of equal size, almost as large as the moderately

sized cusp, form the sickle-shaped anterior part. The jaw is

robust, with fairly coarse denticles. Posterior to the slightly

slanting cusp, along the almost straight inner margin, there

are two, or, in a few cases, one intermediate denticle, the

anteriormost one relatively small, the second larger. The

intermediate dentary is not particularly distinguished. All

denticles on the denticulated ridge , including the in terme

diate ones, slant slightly towards the poste ri or. The number

of denticles varies from 6 to 9 , more or less depending on

the size of the jaw. The shank is about 0.6 of the jaw length,

coarse, almost straight with almost paralleI inner and outer

margins and with a blunt end. The bight is deep with an

acute angle. The ramus is long, of ten straight, and fairly

narrow, tapering distally and ending in a very pointed tip.

Many speeimens have a very vague sinus on the anterior

outer margin at the base of the ramus.

VentraI side : The slightly enclosed, crescent-shaped myer

coele opening represents �3-3f4 of the jaw length . It ex

tends to the anterior along the inn er side and is sur

rounded by a normal rim slightly widened on its outermost

part of the ramus. Left MIl, dorsal side : Length 0 .31-1 .90 mm, width about

. 0 .6 of the length . A double cusp with parts of equal size,

slanting slightly in the posterior direction, forms the anter

iormost, sickle-shaped margin . It is followed by 6-7 inter

mediate denticles of moderate, almost equal size, and

about 8-1 0 sIightly slanting denticles on the straight den

ticulated ridge . The denticle size decreases towards the

posterior end. The shank occupies about half of the jaw

length. The inner margin of the prominent, laterally pro

jecting, inner wing represents about half of the jaw length.

The posterior part of the inner wing is smoothly rounded

and its anterior �3 is paralleI to the denticulated ridge. The

inner and outer margins of the shank are almost paralleI,

and the shank en ds fairly bluntly. The bight is deep with an

acute bight angle. The ramus is long, tapering and pointed


with a minor sinus i n the anteriormost part o f the outer

margin.

VentraI side: The slightly enclosed myocoele opening is

almost a mirror image of the right MIl except that the

opening is less extended anteriorly and thus represents

1J2-�3 of the jaw length.

Remarks. - Normally this speeies is represented by large

jaws. It is found from the Wenlock to Late Ludlow, partic

ularly in the limestone beds on Gotland; it is fairly common

in erratic boulders from the Baltic area (Kielan:J aworowska

1 966) . The morphology becomes more and more conspic

uous with time, with a progressively more developed, prom

inent, needle-like, pointed ramus of the MIl and a very

angular basal portion of the MI, of ten with a spur.

In some localities from the uppermost Slite Beds and the

Halla Beds, an early K. ( K. ) polanensis variety, gandarve

(after the locality Gandarve l ) is found. Typical speeimens

occur in the Middle to Late Wenlock strata; Fardume l , Gandarve l (Fig. 42C) , and Gothemshammar 7 (Fig. 42D ) .

The variety is also found from the Hemse Beds (e .g . Sjaust

rehammar l ) , and related forms have be en encountered

from the Hamra Beds. The gandarve variety seems to occur

partly in paralleI with the Kettnentes (K.) polanensis lineage

on Gotland. The most conspicuous character of the gand

arve variety is the even-sized denticles on the right MIl,

where no cusp can be identified. The inner margin of the

MI is almost straight and the spur is not pronounced. The

basal portion is extremely angled, almost like a square.

Another variety, K. ( K. ) polanensis var. sjaustre, is de

scribed separately below.

Discussion. - The type specimen consists of an apparatus

with the four main elements, including the MIll. It is a very

small jaw apparatus, stored in glycerin, and is thus difficult

to study. To the drawing of the main part of the apparatus

(Fig. 41 ) may be added the following observations. The

right MI has a distinet spur and a fused basal plate . A fused

basal plate seems to be more common among the smaller

right MI ' s than among larger ones within the speeies. Thus,

the fused basal plate may simply reflect the small size of the

speeimen. The outer posterior part of the basal plate is

strongly bent down, which changes the outline of the pos

terior part somewhat. The shank is slightly pointed and almost triangular, the inner wing is straight. The denticles

are fairly large . The left MI has an angular basal portion, an

almost reetangular inner wing, a small spur and large denticles. The right MIl has no cusp, a needle-shaped ramus,

and no distinet sinus. The right MIl has a double cusp, five

intermediate denticles and an almost straight anterior

outer margin .

Kettnentes ( K. ) polanensis dominates the paulinitid fauna

with large jaws (normal 1-1 .5 mm in length) at Glasskar

1-3 and is characteristic in the lowermost Burgsvik Beds of

Ludlow age .

Comparisan. - The species is most easily identified by the

needle-shaped ramus of the MIl and the denticle form ula.

The formula with two pre-cuspidal denticles is not unusual

in the right MIl, but their large, equal size makes them

more easy to distinguish. Of the MI, the almost quadratic

82 Claes F. Bergman

basal portion with the spurs are the most conspicuous

morphological details ( see also K. (K.) burgensis) , although

the shape of the basal portions has a tendency to change

somewhat with time . K. (K.) bankvaetensis has the same

dentition formula but differs in shape and size, see com par

ison of K. ( K. ) bankvaetensis.

The left MIl, no. 738/56, described by Mannil & Zaslaw

skaya ( 1 985a) from the Wenlock Srednij sequence is very

similar to K. ( K. ) polonensis from Gotland.

Kettnerites ( K. ) polonensis var. sjaustre Fig. 46A

Synonymy. - Kettnerites ( K. ) polonensis var. sjaustre - Berg

man, pp. 71-73, Fig. 46A.

Material. - l right MI, l left MI, 2 right MIl, 4 left MIl.

Occurrence. - Fig. 4; Ludlow, Hemse Beds unit d, at present

known only from the locality Sjaustrehammar l .

Diagnosis. - Right MI: Inner margin slightly convex with

about 22 fairly large denticles of almost equal size. Shank

ends sharply. Flange thick-walled, angular. Basal portion

wide, basal angle about 30° .

Left MI: About 28 denticles on the convex inner margin,

denticles decrease in size posteriorly, ending as small

knobs. Inner wing almost rhombic. Wide basal portion,

basal angle about 30° .

Right MIl: Two large pre-cuspidal denticles followed by

a somewhat larger cusp. Two distinct intermediate denti

cles of unequal size. Large shank with almost parallei sides,

ending bluntly. Ramus long, narrow with pointed extrem

ity.

Left MIl: Double cusp with the parts of equal size, distinct

intermediate dentary, shank with almost parallei sides and

ram us long and slender.

Description. - Right MI, dorsal side: Length 1 .47 mm, width

slightly more than a quarter of length . The jaw tapers from the basal part towards the anterior end. The inner margin is sligh tly convex, covered for just over half of the jaw length by 23 fairly large , slightly spaced denticles of equal size. The

posterior part of the undenticulated ridge is slightly bent

outward and is smooth, fairly thick, forming the very sharp

posterior tip of the wide, almost triangular shank. The

inner wing is strongly downfolded, of normal size, 0 .24 of

the jaw length, paralleI to the undenticulated ridge. The

basal furrow is fairly long, its posterior part narrow. The

flange is very thick-walled, its posteriormost part elevated,

the posterior and outer margins forming two nearly right

angles. The anteriormost outer margin of the basal portion

is wide, thick, with a deep, short gro ove separating it from

the inner part of the jaw. The posterior outer part of the

flange is slightly downfolded. The basal portion is wide; the

basal angle about 30° . The outer margin, anterior to the

basal portion, is almost straight and passes gradually into

the large , sickle-shaped falx.

Ventrai side: Not studied.

Left MI, dorsal side: Length 0.95 and 1 .45 mm, width

slightly more than a quarter of length. The jaw tapers from


the anterior half to the large fang. The inner margin is

convex from the undenticulated ridge to the anteriormost

part of the falx and is covered by fairly large denticles

decreasing in size towards the posterior end, ending as

small crenulations close to the anterior margin of the inner

wing. The denticulation covers 0.6 of the jaw length. The

undenticulated ridge is wide and fairly low. The inner wing

is of normal size, 0 .26 of the jaw length, almost rhombic;

the inner margin is parallei to the undenticulated ridge.

Basal furrow short, deep with parallei sides. The posterior

margin is wide, almost oblique to the extension of the inner

margin. The outer margin of the basal portion is short, 0 .24

of the jaw length, and is dominated by the narrow, convex,

outer wing, forrned by the ligament rim. The outer face of

the basal portion is large and convex. The basal portion is

almost rectangular, wide and short; the basal angle about

30° . The outer margin anterior to the basal portion is

straight and passes gradually into the smooth, large, sickle

shaped falx.

Ventrai side : not seen.

Right MIl, dorsal side : Length 0 .48-0.88 mm, width

about half the length. Two large pre-cuspidal denticles of

equal size form the anteriormost, acute-angled margin.

The cusp is of moderate size, slanting slightly in the poste

rior direction. Two denticles of unequal size form the

distinct intermediate dentary. It is followed by nine poste

riorly slanting denticles decreasing in size towards the pos

terior on the almost straight denticulated ridge . The den

ticulation ends well before reaching the blunt end of the

shank. The shank occupies about half of the jaw length; its

almost straight sides are almost paralleI. The bight is deep

with an acute bight angle. The ramus is long and narrow

with a pointed extremity. The anterior outer margin is

straight.

Ventrai side : Not studied.

Left MIl, dorsal side: Length 0 .31-1 . 1 9 mm, width about

half the length . A double cusp with parts of equal size forms

the anteriormost margin. A distinct intermediate dentary is

composed of six relatively small denticles and a larger one

at the transition to the denticulated ridge . Nine fairly large denticles, slanting posteriorly, decreasing slightly in size towards the posterior, form the almost straight, denticu

lated ridge. The shank, with almost parallei sides, occupies

about half of the jaw length . The inner wing is almost

triangular, representing about half of the jaw length, its anterior part widest. The bight is deep with an acute bight

angle. The ram us is long, narrow, with a sinus in its ante

rior, outer margin.

VentraI side: The slightly enclosed, crescent-shaped myo

coele opening represents about 1/2�3 of the jaw length .

The opening is only somewhat extended to the anterior

along the inner side of the jaw. The ligament rim is of

normal size, slightly widened at the outer side, along the

ramus.

Discussion. - Jaws very similar to K. ( K. ) polonensis var.

sjaustre or even identical with this variety have been re

corded from the Polish erratic boulder 410/3 ( the boulder

described by Kielan-jaworowska 1 966) .

FOSSILS AND STRATA 25 ( 1989) Silunan pauli nitid polychaetes from Gotland 83

Fig. 46. All specimens in dorsal view, exeept B5. DA. Kettnerites (K) polonensis var . sjaustre, Sjaustrehammar l , Hemse Beds, unit d, 82-19LJ; Al left MI, LO 581 6:7, x80; A2 right MI, LO 581 6:8, x80; A3 Ieft MIl, LO 581 6:5, x80; A4 right MIl, LO 581 6:6, x80 . DB. K ( K ) sp. A, Likmide 2, Hemse Beds, Hemse Marl SE, 82-28LJ; B1 left MI, LO 5800: 1 , x60; B2 right MI, LO 5800:2, x 1 20; B3 left MIl, LO 5800:3, x60; B4 right MIl, LO 5800:4, x60; B5 basal plate , dorsal-lateral view, note the denticles on the inner side, LO 5800:5, x210 ; B6 same spe eimen as B5, dorsal view, LO 5800:5, x210 .

84 Claes F. Bergman

Remarks. - The variety is named after the locality Sjaustre

hammar l from the Hemse Beds, unit d, of Ludlow age . A

characteristic element is the left MI (Fig. 46A: 1 , LO

581 6:7) .

K. (K.) polonensis var. sjaustre is a rare form on Gotland,

found only at Sjaustrehammar 1. I have placed it as a variety

wi thin K. (K.) polonensis because of the fairly similar den ti c

ulation and ramus of the MIl and the slight similarity of

denticulation and habitus of the basal portion of the right

MI.

Comparison. - The basal portion of the MI is wider than the

corresponding part in K. ( K. ) polonensis.

Kettnerites ( K. ) Sp . A Fig. 46B.

Synonymy. - Kettnerites (K. ) sp. A - Bergman, pp. 69-7 1 , Fig.

46B.

Material. - ane jaw of each, right MI, left MI, right MIl, left

MIl.

Occurrenæ. - Fig. 4; Early Ludlow, known only from the

locality Likmide 2. The sample may be the only one at

present available from this horizon (personal communica

tion, LennartJeppsson) .

Diagnosis. - Right MI: Jaw tapers anteriorly, denticles fairly

large, slanting, largest in the middle part of the convex,

paucidentate ridge . Sharply pointed wide shank; thick

walled flange . Basal angle about 45' .

Left MI : Denticles along convex inner margin varying

from large to small, decreasing in size posteriorly. Inner

wing elongated, rounded, extended posteriorly. Basal

angle about 45' .

Right MIl: Two very large pre-cuspidal denticles, inter

mediate denticles lacking, denticulated ridge convex later

ally, ramus slightly convexo-concave, slender and pointed.

Left MIl: Double cusp with anteriormost cusp slightly

larger, prominent, almost triangular inner wing, ramus long and needle-shaped.

Description. - Right MI, dorsal side : Length 1 .52 mm, width

about V4 of length, or slightly more. The jaw tapers anteriorly, ending in a large fang. The inner margin is slightly

convex, covered by about 1 7 slanting denticles, widely

spaced in the anterior part and largest on the middle part

of the margin. The distance between denticles decreases

posteriorly; on the posterior, middle part the denticles are

closely spaced; the denticulation ends as a few knobs. The

undenticulated ridge is fairly wide, rounded, sigmoidal ,

forming the posteriormost, sharp, pointed tip of the wide

shank. The inner wing is downfolded, fairly short, and the

inner margin is parallei to the undenticulated ridge . The

basal furrow is wide and vaguely defined. The flange is

thick-walled, elevated posteriorly. The outer side of the

shank and the posterior side of the flange form a right

angle , and so do the posterior side of the flange and the

outer side of the flange. The anteriormost outer margin of

the basal portion to the right of the deep and narrow

groove is thick and wide.

FOSSILS AND STRATA 25 ( 1 989 )

The basal plate is not recorded in place - a basal plate

possibly belonging to this variety is almost rectangular with

a deep downfolded outer part. It is denticulated with a

small cusp-like denticle followed by five denticles arranged

parallei with the undenticulated ridge .

The basal portion is wide, the basal angle 40' . The outer

margin in front of the basal portion is almost straight, with

a minor concavity in its middle part, ending with a well-de

fined, sickle-shaped falx.

Ventrai side: not studied.

Left MI, dorsal side : Length 1 .06 mm, width about V3 of

length . The jaw tapers anteriorly, ending in a prominent

fang which is moderately bent upwards in relation to sur

face. The inner margin is slightly sigmoidal with large,

widely spaced denticles in the anterior position. The den

ticulation, 0 .6 of the jaw length, consists of about 17 denti

cles decreasing in size posteriorly and ending in a crenula

tion before reaching the inner wing. The transition into the

undenticulated ridge is smooth; the ridge is slightly arched,

fairly short and wide, parallei to the inner margin. The

inner wing is elongatedly rounded, extending posteriorly,

fairly short, about 0 .22 of the jaw length, widest in the

posterior middle half. The basal furrow is short, relatively

wide and deep. The outer face and margin of the basal

portion is short and narrow, the ligament rim is visible on

anterior margin . The basal angle is about 40' . The lower

middle part of the outer margin anterior to the basal

portion has a small convexity. The falx is large and smooth

rounded.

Ventrai side : Not studied.

Right MIl, dorsal side : Length 1 . 34 mm, width more than

half the length . There are two very large pre-cuspidal den

ticles of equal size, the anterior one forming the almost

pointed anterior end of the jaw. They are followed by nine

very large denticles that slant slightly posteriorly; the

anteriormost of these may be the cusp . Intermediate denti

cles are lacking. The posteriormost two denticles are very

small . The inner margin is convex, the outer slightly con

cave, making the large shank taper slightly towards the

blunt posterior end. The bight is deep with an acute bight angle . The ramus is large with a pointed extremity, the outer margin is convex, the inner concave . A small sinus is

present at the anterior outer margin, at the base of the

ramus. Ventrai side: Not studied.

Left MIl, dorsal side : Length 1 .33 mm, width more than

half the length . The anteriormost part of the double cusp

is slightly larger and forms the sickle-shaped anterior mar

gin. The intermediate dentary with three small denticles of

equal size passes into the denticulated ridge through two

denticles of increasing size. These are followed by about

eight denticles, decreasing in size to the posterior and

slanting strongly towards the outer, posterior side . The

almost straight shank occupies about half of the jaw length,

with a blunt extremity. The nearly triangular, smoothly

rounded, laterally extended inner wing is very prominent,

occupying about half the jaw length, with its widest part

anteriorly. The bight is fairly deep with an acute bight

angle. The ramus is long and slender with a pointed ex-


tremity. The anterior outer margin of the jaw is almost

straight with a minor sinus at the anteriormost part of the

ramus.

VentraI side : Not studied.

Remarks. - The form is placed under open nomenclature

due to its rareness. I believe that K. (K.) sp. A is closely

related to K. (K. ) polonensis because of the dentition and

the form of the ramus of the MIl, and the form of the basal

portion of the MI, excluding the possibly correctly identi

fied denticulated basal plate. The affinities of the taxon are

not yet fully understood, as only one reconstructed appara

tus, composed of five elements, is at hand.

A characteristic element is the right MI (Fig. 46B:2 , LO

5800:2 ) .

Kettnerites ( K. ) versabilis Bergman 1 987 Figs. I 4K-N, I SG, 47

Synonymy. - Kettnerites (K.) versabilis n.sp. - Bergman, pp.

73-76, Fig. 1 4K-N, 18 , 47.

Derivation of name. - Latin versabilis, changeable, referring

to the variable denticulation of the MI and MIl .

Holotype. - LO 5361 :4, right MI, Fig. 47E:2 .

Type locality. - Buske 1 .

Type stratum. - Lower Visby Beds, unit e .

Material. - Fig. 2 ; about 5 1 right MI, 52 left MI , 42 right MIl,

and 40 left MIl .

Occurrence. - Figs. 4 and 6; Upper LIandovery. Lower Visby

Beds, unit d and e, and Upper Visby Beds: Buske l , Gustavs

vik l , Håftingsklint l , Ireviken l , Ireviken 3, Nygårdsbåck

profilen l , Nyhamn l and 4, Lickershamn 2, Vattenfallspro

filen 1 . The species has been encountered only from some

samples from outside the Upper Visby Beds, unit e.

Diagnosis. - Right MI: Jaw tapering anteriorly, cross-section

almost rounded, long slender pointed fang. Anterior den

ticles large; dentary paucidentate; wide, almost pentagonal

basal portion with basal plate fused.

Left MI: Jaw tapering anteriorly, paucidentate dentary

along the anterior part of the straight inner margin; wide almost pentagonal basal portion.

Right MIl: Large single cusp bent strongly upwards in

relation to dorsal surface, intermediate denticles may

occur on the falcal arch.

Left MIl : Large, slender, single cusp , wide-spaced inter

mediate denticles, fairly slender ramus with pronounced sinus on the anterior outer margin.


width about 1f4 of length . The dorsal surface is convex, the

jaw tapering slowly and unevenly towards the anterior. The

inner margin is almost straight, denticulated along its en

tire length in larger jaws; in some jaws also along part of the

falcal arch. In smaller jaws the dentary ends before reach

ing the undenticulated ridge . The anteriormost 1-3 denti

cles in the falcal arch are normally fairly small and followed


by about 5-7 fairly large to very large denticles which slant

slightly towards the posterior. In larger jaws these anterior

denticles are more or less widely spaced to paucidentate .

The denticles of the smaller jaws are only slightly spaeed.

The forms with these different types of dentary are fairly

distinct and their sizes overlap .. The denticulation ends at

the posteriormost part of the denticulated ridge, posterior

or close to the transition into the small undenticulated

ridge at the same level as the inner wing. The undentic

ulated ridge ends before reaching the posterior margin.

The inner wing is almost triangular, its anterior part widest.

The basal portion is large and almost pentagonal. The

basal furrow is fairly deep and of normal length . A fairly

distinet furrow runs along the outer margin of the basal

portion anterior to the basal plate , separating the charac

teristic wide outer margin from the inner part of the basal

portion .

The basal plate varies from almost square in smaller

speeimens to rectangular in larger ones, with height about

half the length of the plate and widely spaced denticles.

The corners of the basal plate form almost right angles, and

the outer margin is deeply downfolded. The basal angle is

about 25-30° .

The outer margin anterior to the basal portion is

rounded longitudinally with a more or less pronounced

posterior concavity. The anterior outer margin along the

fang is smoothly sickle-shaped and ends in a large fang with

a prominent cutting-edge .

Ventrai side : The strongly enclosed myocoele opening is

about 1/3 of the jaw length . The crescent-shaped anterior

margin is surrounded by a fairly narrow and somewhat

sunken ligament rim in the anterior inn er part. Along the

anterior outer part, the rim is bent over towards the dorsal

side and forms a sharp ridge along the inner margin of the

inner wing.

Left MI, dorsal side : Length 0.43-1 . 1 7 mm, width slightly

less than 1/4 of length. The fang is large and pointed, and

the inner margin straight to almost straight. The dentary

varies from normal-spaced in severaI of the smaller jaws to

widely-spaced and paucidentate in some of the larger jaws.

The small anteriormost denticle is followed by 1-2 slightly

larger denticles, which in turn are followed by 1 2-16 den

ticles of uneven size, varying from large to very large . Usually the latter denticles slant slightly and decrease in size towards the posterior. The dentary ends as small knobs

well before reaching the inner wing (Fig. 47E: 1 ) except in

some of the larger jaws with wide-spaced denticles, where it

continues posteriorly to the undenticulated ridge (Fig.

47F: 1 ) . The transition to the undenticulated ridge is

smooth, except in larger jaws with a wide-spaced dentary.

Here, the denticulation ends at the anteriormost part of

the undenticulated ridge, and the ridge is elevated and

bent slightly to the outer side . The basal portion is almost

pentagonal, wide, and has fairly straight margins. The fairly

short and wide basal furrow is paralleI to the outer margin

of the basal portion. The inner margin of the inner wing is

paralleI to the undenticulated ridge . The inn er wing is

almost rectangular, about 1f4-l/5 of the jaw length . A more

or less distinct furrow runs along the characteristic wide



outer margin of the basal portion, ending be fore reaching

the posterior margin. The basal angle is 20-30° . The outer

margin, anterior to the basal portion, has a slightly convex

end. The smoothly sickle-shaped fang has a prominent

cutting edge .

Ventrai side: The strongly enclosed myocoele opening is

almost a mirror image of that of the right MI, except that

it represents about one quarter of the jaw length .

Right MIl, dorsal side: Length 0 .25-0.74 mm, width

about half the length. The large, sickle-shaped, single cusp

is bent strongly upwards in relation to the dorsal side . The

1-3 small denticles on the fa1cal arch represent the inter

mediate dentary on many of the jaws. The transition to the

post-cuspidal dentary is gradual . The size of the denticles

increases towards the posterior, reaching maximum size at

the upper middle part of the dentary. The approximately 8

fairly large denticles slant and decrease in size towards the

posterior, ending in a minor denticle. The shank repre

sents about half the jaw length or slightly more . The outer

margin of the shank is almost straight (Fig. 47A-C; the

bight is of normal size, and the bight angle varies from 60°

to almost 90°. The ramus is slender and slightly pointed,

with a well developed sinus on the anterior outer margin .

Ventrai side : The crescent-shaped, slightly enclosed myc>

coele opening, �3_3/4 of the jaw length, has an anterior

inner bulge and is surrounded by a fairly wide ligament

rim.

Left MIl, · dorsal side : Length 0.43-0.93 mm, width

slightly less than half the length . The sickle-shaped, single

cusp, is large, slender and strongly bent upwards in relation

to the dorsal surface . The 1-3widely spaced intermediate

denticles are followed by 9-1 1 slightly spaced post-cuspidal

ones on the denticulated ridge . The cutting edge is very

distinct. The shank occupies about half the jaw length and

tapers to the posterior. The inn er wing is fairly narrow and

rounded, extending over slightly more than half the jaw

length . The outer margin of the shank is almost straight.

The bight is fairly deep; the bight angle varies from 60° to

almost 90° . The ramus is slender, slightly pointed, with a

pronounced smooth sinus in the anterior outer margin,

forming a very prominent sigmoidal curvature .

Ventrai side: The slightly enclosed myocoele opening is

somewhat more than half the jaw length . In other respects it is almost a mirror image of the right MIl.

Remarks. - It cannot be excluded that with more material it

will be possible to further subdivide this variable taxon.

Three groups based on the MI ' s can be distinguished:

•

Fig. 4 7. Kettnentes ( K. ) versabilis including forms A, B, and C. Speeimens F represent form A, those of D and E form B, and those of G form C. Specimens A, B, E and G are from Buske l , Lower Visby Beds, unit e, 79-40Lj, the remaining specimens (C, D and F) are from Håftingsklint l, Upper Visby Beds, 76-1 0CB. All speeimens are in dorsal view, x1 20, except F. DA. Right MIl, LO 5361 : 1 . DB. Right MIl , LO 5361 :2 . D e. Right MIl , LO 5762: 1 . O D. D l left MI, LO 5762:2 ; D2 right MI, LO 5762:3 . DE. El left MI, LO 5361 :3; E2, holotype, right MI, LO 536 1 :4 ; E3 left MIl , LO 5361 :5 ; E4 right MIl, LO 5361 :6. O F. Fl left MI, LO 5762:4, x80; F2 right MI, LO 5762:5, x80. OG. Gl right MI, LO 5361 :7; G2 left MIl, LO 5361 :8 ; G3 right MIl, LO 5361 :9.


(A) A long and slender form, of normal colour (dark

brown) , with paucidentate, large denticles. The holotype

(Fig. 47F:2) is included in this group.

(B) A slender form, of normal colour, with more or less

wide1y spaced, fairly large denticles (Fig. 47E, D ) . (C) A long slender form, black in colour, with a large

paucidentate dentary (Fig. 47G: l ) . This form is more slen

der and has more pronounced paucidentate dentary than

the holotype.

The jaws show severai similar characters: The MI's of all

three types taper even ly towards the anterior, and all the

right MI's have a fused basal plate . The dorsal surface is

fairly convex and the basal portion fairly large to large .

However, the denticulation of MI 's varies considerably, at

least between forms A and B.

The two forms A and B might be different ontogenetic

stages of the same species (e .g . form B juvenile and form A

adult) or, less likely, be sexual dimorphs.

The size-ranges of the elemen ts overlap to a great extent.

The fairly rare C form is distinguishable by the very unusual

opaque and black colour compared to the brownish colour

that is normal in paulinitid jaws. Due to the limited mate

rial it cannot be resolved whether or not the C form repre

sents another taxon, is a variety of the A form, or is an A

form which has been altered diagenetically.

So far, I have not been able to make a reliable distinction

between the MIl elements associated with the different MI

forms. If, indeed, particular MIl elements are associated

with each of the MI forms listed above, all these MIl types

might not even yet have been encountered in the very

limited material available . However, differences in den ti

tion formulas have been noticed among the MIl elements

(e .g . Fig. 47E:3 and 47G:2) , but with the limited material it

is not possible to appreciate the variation. The MII ' s which

have been linked with the B form of MI (Fig. 47E:3 , 4)

could be associated with the C form of MI (Fig. 47G:2 and

3) and vice versa.

The A and B forms normally occur together and are also

associated with the C form in rich localities . The C form

(Fig. 47G: l ) has so far be en found only in Buske l , Håft

ingsklint l , and Nygårdsbåckprofilen l , from which locali

ties the largest collections of the species (also A and B

forms) derive . About 8 right MI and 5 left MI have been encountered of form C.

Comparisons. - The basal portion of the MI ' s shows a slight

similarity to that of K. ( K. ) polonensis var. sjaustre , but the

MI of the latter has a wider outer margin (outer wing) .

Kettnerites (Aeolus) Bergman 1 987

Synonymy. - 0 1987 Kettnerites (Aeolus) n. subgen. - Berg

man, p. 77.

Derivation of name. - From the Greek mythology; Aeolus was

the ruler of the winds, father of Sisyphus.

Type species. - Kettnerites (Aeolus) sisypki Bergman 1 987

Dtker species. - K. (A.) microdentatus, K. (A.) fjaelensis, K. (A. )

siaelsoeensis.

88 Claes F. Bergman

Diagnosis. - Right MI: Fairly slender jaw, slender fang. Pro

nounced, alm ost rounded, of ten fairly wide basal portion .

Left MI: Slender jaw, basal portion with almost straight

posterior and outer margins, inner wing almost rectangu

lar.

Right MIl: One or two very small to fairly small pre-cuspi

dal denticles. Cusp fairly large, its base swollen .

Left MIl : Fairly large cusp without pre-cuspidal denticles,

except for K. (A. ) sisyphi var. valle which has one.

Remarks. - The denticulation of the left and right MI varies

considerably, from fine, densely-spaced denticles of equal

size to fairly large denticles of unequal size. The denticles

of the left MI are slightly finer than those of the right MI.

The dentition on the inner margin also varies in extent

between the different subspecies of the corresponding ele

ment type. The dentition of the right MI is usually more

extended than on the left MI.

The number, arrangement and shape of the denticles

(especially the pre-cusps and the cusp) of the right MIl and,

to a lesser degree, of the left MIl , varies between the

subspecies. This variation also occurs between populations

of K. (A.) sisyphi sisyphi from the Middle-Late Wenlock.

There is no particular variation of the dentition (form,

extension) , between the same element type (left and right

of MI and MIl) within the populations in a locality (sam

ple) .

Kettnerites (Aeolus) sisyphi Bergman 1987 Figs. 1 2K-O, 1 8H, 20, 48-53

Synonymy. - 0 1 985a Kettnerites aspersus (Hinde 1 880) -

Miinnil & Zaslawskaya, pp. 1 1 7-1 18 , Pl. 1 7 :4e, right MI

(738/55) . 0 1987 Kettnerites (Aeolus) sisyphi n. sp. - Berg

man, p. 77-87, Figs. 1 2K-O, 18 , 20, 48-53.

Derivation of name. - As for the nominal subspecies.

Holotype. - As for the nominal subspecies.

Type locality. - As for the nominal subspecies.

Type stratum. - As for the nominal subspecies.

Subspecies included. - K. (A.) sisyphi sisyphi, K. (A.) sisyphi

klasaardensis, K. (A. ) sisyphi var. valle .

Material. - More than 250 elements of each of the four main

elements.

Occurrence. - Late Llandovery to Late Ludlow. Lower Visby

Beds to Hamra Beds.

Diagnosis. - Right MI: Jaw with almost parallei inner and

outer margins and pronounced basal portion with large,

deep, downfolded flange . Denticles of varying size, smallest

in central part and largest in the posterior or anterior part,

ending abruptly at the undenticulated ridge .

Left MI: Slender jaw, tapering anteriorly. Denticles fairly

small, ending as small crenulations well before reaching

the undenticulated ridge . Almost rectangular inner wing

with inner margin straight and parallei to the undentic

ulated ridge .


Right MIl: Two (except in K. ( K. ) sisyphi klasaardensis

which has one) very small to fairly small, pre-cuspidal den

ticles. Cusp fairly large , more or less swollen in its basal part.

Shank fairly pointed, bight angle almost 90' .

Left MIl : Fairly large , single cusp, except in K. (A. ) sisyphi

var. valle which has one pre-cuspidal denticle. Four to six

intermediate denticles. Shank fairly pointed; bight angle

almost 90' .

Description. - Right MI, dorsal side : Length 0 .21-2 . 1 mm,

width about lf4 of length. The slender jaw, narrowing at the

midd le , ends an teriorly in a pronounced sickle-shaped falx,

and has a distinct cutting edge along the anterior outer

margin . The inner margin is straight, denticulated from

the falcal arch to the undenticulated ridge . The dentition

is fairly dense, except in K. (A. ) sisyphi klasaardensis, where

the anterior denticles are slightly spaced. The size of the

denticles varies considerably in some populations, but also

within a single jaw. Usually they are fairly small at the

middle of the dentary and larger in the posterior or ante

rior part. The anterior denticles are often oriented slightly

forward or obliquely to the length axis of the jaw, and the

posterior ones slant towards the posterior. The denticu

lated ridge ends abruptly with distinct denticles at the

transition to the undenticulated ridge which is fairly high,

narrow and of ten slightly bent outwards at the middle,

ending in the posteriormost part of the shank.

A square basal plate with rounded corners is recorded

from K. (A. ) sisyphi klasaardensis and K. (A. ) sisyphivar. valle.

The basal portion is fairly wide , the basal angle about

1 0-30' . The inner wing is deeply downfolded, fairly narrow

at the posterior, widening anteriorly and ending in a fairly

large, rounded anterior corner. The bight angle is almost

90' . The flange is normal to slightly thickened, wide and

deeply downfolded along the posterior outer margin. A

more or less conspicuous, laterally extended border runs

along the anterior outer margin of the basal portion. The

outer margin anterior to the basal portion and posterior to

the falx has a more or less pronounced concavity. Left MI, dorsal side : Length 0 .35-3 .0 mm, width about

V4 of length . The jaw is slender, normally with a pro

nounced falx and a prominent cutting edge along the

outer margin. Denticle size decreases towards the posterior. The denticulation ends as small knobs or crenulations

in the middle, or slightly posterior to the middle, on the denticulated ridge, well before reaching the inner wing.

The transition to the low, undenticulated ridge is smooth

and gradual . The basal portion is fairly w!de and conspicu

ously skewed rectangular to almost pentagonal. The inner

wing is almost rectangular, angular or rounded, the basal

furrow of normal size. The posterior outer margin of the

basal portion is almost straight, the basal angle about 30' .

The outer margin anterior to the basal portion and poste

rior to the falx is slightly concave.


about half the length. There are two very small to fairly

small pre-cuspidal denticles ofunequal size on the anterior

most part of the large cusp , except in K. (A. ) sisyphi klasaard

ensis which has only one. The almost straight, denticulated

ridge is composed of fairly large to large denticles, slanting


posteriorly, with the largest ones in the middle . The shank

occupies slightly more than half the jaw length; the almost

straight inner and outer margins taper to the posterior.

The bight is fairly deep, the bight angle varies between

acute and 90° . The ramus is fairly narrow, the anterior

outer margin, almost straight.

Left MIl, dorsal side : Length 0. 1 6--0.83 mm, width about

half the length . There is a large, sickle-shaped cusp with a

distinct cutting edge, and no pre-cuspidal dentary, except

in K. (A. ) sisyphi var. valle. The cusp is followed by 4-6 fairly

large intermediate denticles. All the post-cuspidal denticles

vary in number and size. The shank is about half the jaw

length and tapers to the posterior. The inner wing is of

normal size. The bight is deep, the bight angle acute to

almost straight. The ramus is almost triangular to long and

narrow, posteriorly somewhat pointed. The sigmoidal

shape of the anterior outer margin is due to a pronounced

concavity at the base of the ramus.

Discussion. - The difference between the subspecies is most

clearly expressed in the pre-cuspidal dentary of the right

MIl. The nominal subspecies has two very small to small

pre-cusps while those of K. (A. ) sisyphi var. valle are of

normal size. The subspecies K. (A. ) sisyphi klasaardensis has

only one minor pre-cusp.

Comparison. - The left and right MIl of K. (A. ) sisyphi sisyphi

are very similar to the corresponding elements of K. (A. )

microdentatus.

Kettnerites (Aeolus) sisyphi sisyphi Bergman 1987 Figs. 1 2L, M, 0, 1 8H, 20 , 48 , 49

Synonymy. - O 1 987 KeUnerites (Aeolus) sisyphi sisyphi n. ssp. -

Bergman, p. 79-8 1 , Figs. 1 2L, M, O, 18 , 20, 48, 49.

Derivation ofname. - Named in commemoration of Sisyphus

who also had a gre at problem to solve .

Holotype. - LO 5836:2 , right MI, Fig. 48D :2 .

Type locality. - Gandarve l . Type stratum. - Halla Beds.

Material. - Figs. 2 and 3; more than 250 right MI, 250 left

MI, 250 right MIl, 250 left MIl.

Occurrence. - Figs. 4 and 7; Late Llandovery to Ludlow.

Lower Visby Beds to Hemse Beds. Ajmunde l , Ajstudden l , Alby l , AmIings l , Ansarve l , Autsarve l , Bjarges l and 2 ,

Blåha\l l , Broa 2 , Dapps l and 2 , Djupvik l , 2 , and 3 ,

Fardume l , Fåro Skola l , Follingbo 3 , Gandarve l and 2 ,

Gannor 3 , Gardsby l , Gerete l , Gnisvard l , Godrings l , Gothemshammar 6 and 7 , Grondalen l , Grundård 2 ,

Grymlings l , Hiiftingsklint l , Haganas l , Hagur l , Halls

Huk l , Hide l , Horsne 5 , Kiilldar l and 2 , Kullands 2, Lau

Backar l , Lauter l , Lerberget l , Lickershamn 2, Lilla Hall

vards l , Loggarve l and 2, Lukse l , Molner l , Munkebos l , Mulde 2 , Mulde Tegelbruk l , Myrsne l , Nygårds l and 2,

Rangsarve l , Robbjans Kvam 3, Ronnklint l, Saxriv l , Sigvalde 2 , Slitebrottet l and 2 , Snackgardsbaden l , Snau-


valds l , Snoder 2 and 3, Sojvide l , Sproge 4, Stora Myre l , Stora Vikare 2 , Svarven l , Talings l , Tjeldersholm l , Tradgården l , Valby Bodar l , Valleviken l , Vallstena 2 ,

Valve 3, Varsande l , Vasterbjars l , Vattenfallsprofilen l , Vidfalle l .

Diagnosis. - Right MI: Fairly slender jaw, pronounced,

rounded basal portion. Dentieles of normal size, ending

abruptly at the undenticulated ridge . Smallest denticles in

the central part of the dentary and largest at the posterior

end.

Left MI: Slender jaw, tapers anteriorly, en ds in a fairly

small fang. Denticles fairly small, covering about 1J2-�3 of

the margin, ending well before reaching the smooth tran

sition to the undenticulated ridge .

Right MIl : Two very small to small pre-cuspidal denticles.

Cusp fairly large, slightly swollen at the base . Shank fairly

pointed, bight angle almost right.

Left MIl : Single, fairly large cusp . Four to six intermedi

ate denticles. Shank slender, fairly pointed; bight angle

almost right.


width about 1J4 of length. The slender jaw, narrowing at the

middle , en ds anteriorly in a pronounced sickle-shaped

fang with a distinct cutting edge along the anterior outer

margin. The inner margin is straight, denticulated from

the fa\cal arch to the undenticulated ridge . The dentition

varies considerably from specimens with fairly small denti

eies in the Early Wenlock and the Early Ludlow to speci

mens with larger dentieles in the Middle Late Wenlock,

particularly those from the Halla and Mulde Beds. The

dentition of specimens from this stratigraphic interval is

also more diversified, with fairly small denticles at the

middle of the dentary and the large st at the posterior end.

The anterior dentieles are oriented slightly forward or

obliquely to the length axis of the jaw, and the posterior

ones slant towards the posterior. The denticulated ridge

ends abruptly with distinct denticles at the transition to the

undenticulated ridge . The undenticulated ridge is fairly

high, narrow and of ten slightly bent outwards at the mid

dIe, ending in the posteriormost part of the shank.

The basal plate is not recorded. The basal portion is fairly wide with rounded corners.

The basal angle is about 1 0-30° . The inner wing is about 1J4 of the jaw length . Its posterior part is fairly narrow,

widening in the anterior and ending in a fairly large and

conspicuous, rounded anterior corner. The bight angle is

almost 90° . The flange is normal to slightly thickened, wide ,

deeply downfolded along the outer side and the posterior

outer corner, and with a ligament rim along the outer

margin of the basal portion . The outer margin, anterior to

the basal portion and posterior to the falx, has a more or

less pronounced concavity.

Ventrai side : The crescent-shaped, strongly enclosed

myocoe!e opening, about 1/3 of the jaw length, is bounded

to the anterior and along the inner margin by a somewhat

sunk ligament rim, forming a narrow ridge along the inner

wing. On the outer side the rim is bent over and is visible

from the dorsal side .


FOSSILS AND STRATA 25 ( 1 989) Silurian paulinitid polychaetes from Gotland 9 1

Fig. 49. Kettnerites (A.) sisyphi sisyphi. A and B form main part o f ajaw apparatus o f fused elements from Arnlings l , Hemse Beds, Hemse Mari, NW part, 75-73CB, x 1 20. DAL Left MI and MIl, stereopair, dorsal view, LO 5828:3; oA2. Lateral view of the same speeimens as Al . DB. Right MI, MIl and MIV, stereopair, dorsal view, LO 5826: 1 .

Left MI, dorsal side: Length 0 .36-0.83 mm, width about

V4 of length . The slender jaw, with a sickle-shaped falx,

ends in a fairly small fang with a prominent cutting edge.

The large st denticles are at the anteriormost part of the

denticulated ridge . They are perpendicular to the exten

sion of the jaw on the almost straight inner margin. The

denticles decrease in size to the posterior, ending as small

knobs or crenulations in the middle of the denticulated

ridge . The posterior denticles are perpendicular to the

extension of the jaw or slant slightly towards the posterior.

The transition to the low, undenticulated ridge is smooth

and gradual. The basal portion is trapezoidal and fairly

wide. The inner wing is almost rectangular and rounded,

the basal furrow of normal size and wide. The posterior

outer margin of the basal portion is almost straight; the

basal angle about 30° . The outer margin between the basal

portion and falx is slightly concave.


Fig. 48. Kettnerites (A.) sisyphi sisyphi. All speeimens in dorsal view, x1 20. DA. Snoder 2, Hemse Beds, Hemse Mari, NW part, 82-14CB; Al left MI, LO 5809 : 1 ; A2 right MI, LO 5809:2 . DB . Vårsånde l , Mulde Beds, lowermost part, 75-22CB; Bl left MI, L O 5793:5; B2 right MI, LO 5793:6; B3 left MIl, LO 5793:7; B4 right MIl, LO 5793:8. De. Dåpps 2, Mulde Beds, upper part, 82-37CB; Cl left MI, LO 5786 : 1 ; C2 right MI, LO 5786:2 ; C3 left MIl, LO 5786:3; C4 right MIl, LO 5786:4. OD. Gandarve l, Halla Beds, 71-8 1LJ; Dl left MI, LO 5836: 1 ; D2, holotype, right MI, LO 5836:2; D3 right MIl, LO 5839: 1 . DE. Robbjåns Kvam 3, Mulde Beds, lower part, 71-1 38LJ; E l left MI, LO 5775 : 1 ; E2 right MI, LO 5775:2 . OF. Vallstena 2, Slite Beds, Pentamerus gothlandicus Beds or slightly older, 77-2CB; Fl left MI, LO 5777 : 1 ; F2 right MI, LO 5777:2 . O G. Vårsånde l, Mulde Beds, lower part, 75-23CB; Gl left MI, LO 5790: 1 ; G2 right MI, LO 5790:2 . OH. Slitebrottet 2, Slite Beds, Slite Mari, 8�31LJ; Hl left MI, LO 5773:3; H2 right MI, LO 5773:4. Ol. Right MI, LO 5362 :3, Buske l, Lower Visby Beds, unit e,

79-40L]. O]. Right MI, LO 5363 : 1 , Lickershamn 2 , Lower Visby Beds, unit f, 73-53L].

lA-V3 of the jaw length, in other respects it is very similar

to the one of the right MI.


about half the length . Two small, pre-cuspidal denticles of

unequal size project from the anteriormost part of the

large cusp. The cusp is oriented alm ost perpendicular to

the length axis of the almost straight denticulated ridge,

which is composed of 2-3 minor denticles followed by 6-8

larger on es, slanting towards the posterior, with the largest

denticles at the middle . The shank occupies slightly more

than half the jaw length, the almost straight inner and

outer margins taper to the posterior. The bight is fairly

deep, bight angle 60° to almost 90° . The ram us is fairly

narrow, the anterior outer margin almost straight. The jaw

has a triangular appearance.

Ventrai side : The slightly enclosed myocoele opening,

�3 of the jaw length or somewhat less, is bounded by a fairly

wide and somewhat elevated ligament rim to the anterior

and along the inner side . The almost straight anterior and

inner margins form a corner which protrudes slightly ante

riorly.

Left MIl, dorsal side : Length 0.22-0.50 mm, width approximately half the length . The large , sickle-shaped, cusp

with a distinct cutting edge, is oriented perpendicular to the denticulated ridge and is followed by 4-6 fairly large

intermediate denticles. The denticulated ridge is slightly

convex with usually 8-9 posteriorly slanting denticles, the

largest in the posterior half of the denticulated ridge . The

shank tapers to the posterior and occupies about half the

jaw length, ending fairly pointed and somewhat bent out

wards. The inn er wing is of normal size. The bight is deep,

the bight angle 60° to almost 90° . The ramus is long and

narrow, posteriorly somewhat pointed. The anterior outer

margin is sigmoidal because of a pronounced concavity at

the base of the ramus.


V2-�3 of the jaw length . The almost straight anterior and

inner margins form an almost right angle . The rim is

normal and becomes wider at the outer part of the ram us.

92 Claes F. Bergman

Kettnerites (Aeolus) sisyphi var. valle Figs. 1 2K, 1 8H:3, 50

Material. - Figs . 2 and 3; more than 59 right MI, 57 left MI,

55 right MIl, 35 left MIl.

Occurrence. - Figs . 4 and 7; Middle Wenlock. Slite Beds

Pentamerus gothlandicus Beds, Slite Siltstone , Halla Beds,

unit b. Gandarve l , Gerumskanalen l, Gothemshammar 8,

Haganas l, Mulde Tegelbruk l , Klinteenklaven l , M6llbos

l, Svarvare l and 3 , Valbybodar l, Valle l and 2.

Diagnosis. - Right MI: Inner and outer margin almost

straight and parallei, denticles fairly large and of almost

equal size, shank ending bluntly.

Left MI: Inner margin almost straight, outer margin

convex. Denticles decrease in size posteriorly. Basal portion

angular, large inner wing almost rectangular, parallei with

undenticulated ridge.

Right MIl: Two pre-cuspidal denticles of normal size on

a sickle-shaped anterior margin. Anterior margin slightly

angular in the smaller specimens. Ramus fairly small and

narrow.

Left MIl: One distinct pre-cuspidal denticle; ramus fairly

small and narrow.

Description. - Right MI, dorsal side : Length 0.35 to about 2 . 1 mm, width about V4 o f length . The anterior part o f the

dorsal surface is almost flat, the posterior part convex. The

inner and outer margins are almost straight and almost

paralleI. The fang is narrow and bent upwards in relation

to the surface . The denticulation of the inner margin cov

ers about 0 .6-0.7 of the jaw length . The approximately 30

denticles are of almost equal size, though the posterior

central and posteriormost ones are somewhat smaller. The

anterior 5 or so denticles are slightly spaced and directed

forward or obliquely with regard to the direction of the

inner margin. The following denticles are closely spaced

and slightly slanting. The denticulation ends posterior to the anterior end of the inner wing with some very small denticles. The high, smooth and narrow undenticulated

ridge is bent outward in the centre, without reaching the

posterior margin. The shank is wide with a blunt end. The

bight angle is acute , close to 70° . The inner wing is of

normal size, almost rectangular, deeply downfolded; the inner margin is parallei to the undenticulated ridge . The

basal furrow is deep and long. The flange is thin-walled and

angular, its outer part deeply downfolded and extending

anteriorly. The anterior part of the basal portion is folded

horizontally, forming a rounded outer wing. The basal

angle is 25-30° , with the inner and outer margins of the

basal portion more or less paralleI.

The basal plate is almost square .

The outer margin of the fal x is somewhat flattened on its

outer anterior part. Along the outer margin a cutting edge

is very distinct for more than half of the jaw length .


myocoele opening, about lf3 of the jaw length, is bounded

in the anterior and along the inner margin by a fairly high

ligament rim which forms a sharp ridge along the inner


wing. Along the outer side , the margin, including the rim,

is bent over and visible from the dorsal side .

Left MI, dorsal side : Length 0 .38-3 .0 mm, width about

V4 of length. The dorsal surface is flat to concave along the

outer margin. The anterior part of the fang is pointed and

narrow. The inner margin is almost straight, denticulated

from the anteriormost part of the fakal arch, for about �3

of the length of the inner margin. The denticles are of

medium size, decreasing toward the posterior end. There

is a smooth transition to the long, smooth and narrow

undenticulated ridge . The inner wing is large, rectangular,

its posterior outer corner prominent, and its outer margin

parallei to the undenticulated ridge . The basal furrow is

narrow, deep and parallei to the outer margin of the basal

portion. The posterior margin is alm ost straight, sometimes

directed obliquely towards the inner margin . The outer

margin of the basal portion is slightly downfolded, wide in

the anterior and forming an outer wing bordered by a

straight ligament rim. The basal portion is almost pentago

nal and angular. The basal angle is 25-30° . The outer

margin is strongly convex, somewhat flattened in the ante

rior outer portion and dominated by the cutting edge

which runs along the outer margin from the anteriormost

part of the falx alm ost to the basal portion.


myocoele opening is about 1/3 of the jaw length or slightly

less. The ligament rim is of normal width and slightly lower

than the jaw surface .

Right MIl , dorsal side : Length 0 .25-l .33 mm, width

slightly more than half the length. There are two pre-cuspi

dal denticles of normal size on the sickle-shaped anterior

margin . Smaller specimens may have a somewhat angular

anterior margin . The cusp is of normal size and slants

slightly posteriorly, with a prominent cutting edge on its

anterior basal part. The denticles slant posteriorly and

increases slightly in size from the anterior towards the

centre of the dentary. There are no distinct intermediate

denticles. The number of denticles is 1 0-16 , more or less

correlated with jaw size. The almost straight shank tapers slightly to the posterior and represents slightly more than

half the jaw length. The inner wing is very minute, following the inner margin, widest at the posterior. The bight

angle varies from acute to almost straight. The ramus is

relatively small, particularly in the larger specimens. It is

fairly narrow and pointed with a slight basal sinus on the

outer, anterior margin.


%_3/4 of the jaw length. The ligament rim is fairly narrow,

Fig. 50. Kettnentes (A. ) sisyphi var. valle . I andJ are referred to as K. (A. ) sisyphi var. cf. valle. Valle 2, Slite Beds, Pentamerusgothlandicus Beds, 66-1 45SL. G, H, L, M, N are x80, all other specimens x1 20. All in dorsal view. DA. Left MIl, LO 5770 : 1 . DB. Left MIl, LO 5770:2 . De. Left Mn, LO 577 1 : 2 . OD. Right MIl, LO 5771 :3 . DE . Right MIl , LO 5771 :4. OF. Right MIl , LO 5770:3. O G. Left

MIl, LO 5770:4. OH. Right MIl, LO 5770:5 . D I . Left MI, LO 577 1 :5 . Oj. Right Ml, LO 5771 :6. DK. Left MI, LO 5771 :7 . DL. Right MI, LO 577 1 :8 . D M. Left MI, LO 5771 :9. ON. Right MI, LO 577 1 : 1 0 .


94 Claes F. Bergman

the opening extends in the an terior inner part to form a

protruding corner. The margins of the opening are almost

straight.


half the length or slightly less. There is one distinct pre

cuspidal denticle with a cutting-edge on the anterior part

of the smoothly rounded anterior margin . The cusp is fairly

thin and alm ost perpendicular to the extension of the

denticulated ridge . The intermediate dentary is not partic

ularly distinct because of the large size of the denticles.

Their num ber varies from 4 to 6 and they are of equal size.

The following denticles are almost of equal size with the

exception of the posteriormost on es, and they slant in

posterior direction. The shank tapers posteriorly and rep

resents less than half the jaw length . The inner wing is

small, with a rounded inner margin . The bight angle is

almost 900 • The ramus is pointed and relative ly small, with

a more or less pronounced sinus at the outer margin of the

base .

Ventrai side: The crescent-shaped, slightly enclosed myo

coele opening extends anteriorly along the inner side for

more than half the jaw length . The ligament rim is of

normal width , widening in the outer part of the ramus.

Discussion. - This taxon, except the left MIl, is very close to K. (A.) sisyphi sisyphi. It is only recorded from a few

stratigraphically restricted localities. K. (A. ) sisyphi var. valle

occurs together with the nominal subspecies. Further ma

terial may indicate whether this taxon should be treated as

a species.

Remarks. - A typical element of K. (A . ) sisyphi var. valle is the

right MI, LO 577 1 : 10 , Fig. 1 3N from Valle 2 , Slite Beds, Slite

Mari , Pentamerus gothlandicus Beds.

Kettnerites (Aeolus) sisyphi klasaardensis Bergman 1 987 Figs. 1 2N, 1 8H: l , 20 , 51-53

Synonymy. - O 1987 Kettnerites (Aeolus) sisyphi klasaardensis n. ssp. - Bergman, p. 83-87, Figs. 1 2N, 1 8, 20, 51-53.

Derivation ofname. - After the type locality, Klasård l , where

this species is dominant.

Holotype. - LO 5823 : 13 , right MI, Fig. 5 1 P.

Type locality. - Klasård l , Gotland.

Type stratum. - Hemse Beds, Hemse Mari SE.

Material. - Fig. 3; more than 157 right MI, 131 left MI, 1 33

right, 1 39 left MIl, 23 MIll, 1 7 left and right carriers.

Occurrence. - Figs. 4 and 7; Ludlow, Leintwardinian. Hemse

Beds, Hemse Mari SE part, Eke Beds basal part. Bodudd l and 3, Hallsarve l , Herrvik 2, Kårne 3, Klasård l , Stora

Kruse l , Vaktård 2, 4, and 5, Vastlaus 1 .

Diagnosis. - Right MI: Almost parallei mner and outer

margins; inner margin straight, smallest denticles in mid

die of denticulated ridge . Large angular, almost pentago

nal basal portion with fused, square, basal plate .


Left MI: Straight inner margin; denticles decrease in size

posteriorly, ending as crenulations without reaching the

nearly rectangular inner wing. Basal portion almost pentag

onal , angular.

Right MIl: One minor, pre-cuspidal denticle , the contin

uation ofwhich forms, in relation to the large swollen cusp,

a characteristic keel along the falx.

Left MIl: Prominent cusp with almost straight anterior

margin .

MIll : Concavo-convex element with pronounced cusp,

preceded anteriorly by three denticles.

Right MIV: Concavo-convex element, in lateral view cres

cent-shaped, without cusp, second and third denticles from

the anterior forming the highest part of the jaw.

Left MIV: Mirror image of the right MIV.

Carriers: A pair of left and right elements, chisel-like,

elongated, thin, with pronounced head and shaft with a

longitudinal furrow.

Description. - Right MI, dorsal side : Length 0 .23-0.96 mm, width approximately 0.3 of length. The anterior part of the

fang is slender, bent upward in relation to the almost flat

jaw surface . The inner margin is straight, denticulated from

the anteriormost part of the falcal arch to the undentic

ulated ridge . The number of denticles is 1 8-30, the number

increasing more or less regularly with increasing jaw

length . The first 2-5 anterior denticles are of ten large,

slightly spaced, some directed forward, or more commonly,

perpendicularly to the length axis of the jaw. The middle

denticles are smaller and densely spaced. The posterior

third of the denticles, again larger, of ten slanting, end

abruptly at the undenticulated ridge with a few small dis

tinct knobs on the margin, which of ten makes a curve here .

The ridge is high, narrow and rounded, slightly bent out

wards at the middle, reaching the posterior end of the shank and forming the posteriormost tip of the jaw. The

inner wing is strongly downfolded, almost triangular with a

rounded anterior inner ' corner' . Almost parallel to and

immediately to the right of the undenticulated ridge is the

deep and narrow longitudinal basal furrow, partly covered by the basal plate .

The basal plate is square, undenticulated, and confluent

with MI on its anterior and inner margins. A low ridge runs

along its inner margin, with a small, short furrow immedi-

Fig. 51. Kettnerites (A.) sisyphi klasaardensis. Klasård l , Hemse Beds, Hemse Mari, SE part, 71-1 50L]. A, B, C, D ventral view;J, K, L lateral view; the remaining in dorsal view. bA. Right MI, basal portion, LO 5823 : 1 , x1 20. DB. Left MI, basal portion, LO 5823:2, x1 20. DC. Right MIl , LO 5823:3, x 120 . OD. Left MIl, LO 5823:4, x1 20. DE. Pair of carriers, stereopair, LO 5824: 1 , x230. O F. Left MI, LO 5823:4, x1 20. OG. Right MI, LO 5823:5, x 120 . O H. Left MI, LO 5823:6, x210. DI. Right MI, LO 5823:7, x210 . O]. Carrier, LO 5824:2, x2 10 . DK Pair of carriers, LO 5824:3, x250. D L. Carrier, LO 5824:4, x170 . D M. Left MIl, stereopair, LO 5823:8, x21 0. ON. Right MIl, stereopair, LO 5823:9, x210 . DO. Left MI, stereopair, LO 5823: 1 0, x1 20. D P. Holotype, right MI, stereopair, LO 5823: 1 3, x1 20. DQ. Left Mil, LO 5823 : 1 2, x 1 20. DR. Right

MIl, LO 5823: 1 1 , x120.




Fig. 53. Kettnerites (A . ) sisyphi klasaardensis. All speeimens from the Hemse Beds, Hemse Mari, SE part, except B; all in dorsal view. DA. Vaktård 4, 81 -35LJ; Al left MIl, LO 5822:3, x 1 70; A2 right MIl, LO 5822:4, x 1 20 . D B. Left MIl, LO 5843:3, Kårne 3, Eke Beds, lowermost part, 71 -198LJ, x 1 20 . DC. Basal portion of right MI including the basal plate in place, LO 5815 : 10 , Bodudd 3, 7l - 151LJ, x280. OD. Bodudd 3, 71 - 151LJ; Dl left MIl, LO 5815:9, x1 20; D2 right MIl, LO 5815 :8 , x120 . DE. Precusp and cusp of right MIl, LO 5823 : 17 , Klasård l, 71-150LJ, x830.

Fig. 52. Kettnerites (A.) sisyphi klasaardensis. F-O are in dorsal position. All x 1 20 except C, D and E. DA. Right MI, ventrai view, LO 58 15 :7, Bodudd 3, Hemse Beds, Hemse Mari, SE part, 71-1 5 1 LJ. D B. Left MI, ventraI view, LO 58 15 :6, same sample as A. DC. Left MIV, lateral view, LO 5823: 1 6, Klasård l , Hemse Beds, Hemse Mari SE part, 71-150LJ, x250. OD. Right MIV, dorsal view, LO 5823: 15 , same sample as C, x250. DE. Same sample as C; E l MIll, lateral view, L O 5823: 14, x250; E 2 same speeimen a s E l , dorsal view, x250. OF. Right MI, L O 5843:2, Kårne 3 , Eke Beds, lowermost part, 71-198LJ. O G. Right MI, LO 5822 : 1 , Vaktård 4, Hemse Beds, Hemse Mari, SE part, 81 -35LJ. OH. Right MI, LO 5804: 1 , Hallsarve l, Eke Beds, lowermost part, 69-28LJ. DI . Right MI , LO 5815 :5 , Bodudd 3, Hemse Beds, Hemse Mari, SE part, 7l-1 5 1LJ. DJ. Right MI , LO 581 5:4, same sample as I . DK Left MI, LO 581 5 : 1 , same sample as I. D L. Left MI, LO 5815 :2 , same sample as I. D M. Left MI, LO 5822:2, same sample as G. ON. Left MI, LO 5815 :3, Bodudd 3, same sample as I. D O. Left MI, LO 5843: 1 , same sample as F.

ately to the right of its posterior half. The posterior half is

slightly concave; the outer margin strongly downfolded. Anterior to the basal plate, on the basal portion, the

ligament rim forms a pronounced outer wing. At a distance

about twice the length of the basal plate from the posterior

end, the rim drops steeply, forming a characteristic tip at

the junetion. The boundary between the rim and the rest

of the jaw is smooth. The basal angle is about 20-25° . In

front of the basal portion the outer margin is concave ,

swings out and ends in the sickle-shaped falx. The distinet

outer cutting-edge of the fang continues along the outer

margin for about V3 of the jaw length .

Ventrai side: The strongly enclosed myocoele opening is

approximately V3 of the length of the jaw. The basal plate

is separated from the myocoele opening by a costa repre

senting the anterior and inner margin of the basal plate .

Left MI, dorsal side: Length 0 . 1 5-0.82 mm, width about

l/4_1Jt of length . The inner margin is straight and the outer

margin parallei to the inner margin between the basal

98 Claes F. Bergman

portion and the falx. The fang is narrow and pointed, fairly

strongly bent upwards in relation to surface of the jaw.

There are two distinct cutting edges on the fang, one on the

inner side continuing onto the denticulated ridge , the

other continuing along the outer margin of the falx. The

inner margin, denticulated from the falcal arch, extends

half the length of the jaw, ending I l/2-2 times the length of

the inner wing from the posterior margin. The num ber of

denticles increases more or less with jaw length, from about

12 to 32. The anterior 2-4 denticles are often slightly larger,

directed forward, more or less widely spaced. The denticles

gradually decrease in size posteriorly, ending as crenula

tions on the inner margin . Thus, the number of denticles

is difficult to discern . There is a smooth transition to the

undenticulated ridge , which is low, narrow, slightly bent

outwards medially, tapering posteriorly, almost reaching

the posterior margin as a narrow downbent ridge . The

almost rectangular inner wing of normal size, bent down

and parallei to the inner margin . The basal portion is

angled, irregularly pentagonal, with the slightly concave

posterior margin perpendicular to the long axis of the jaw.

The basal furrow is short and almost open towards the

posterior outer side . The basal angle is about 20-35° . The

outer face of the basal portion is large and smoothly con

vex, downfolded along the posterior outer margin. A more

or less pronounced outer wing runs along the outer margin

of the basal portion. It is separated from the outer face by

a distinct, narrow groove . The wing rises anteriorly, form

ing a peak about V5 of the jaw length from the anteriormost

part of the wing, whence it dips down again .


myocoele opening is about V3 of the jaw length . An almost

flat, fairly broad ligament rim forms a smaller ridge along

the inner margin of the inner wing. On the outer side, the

ligament rim continues on the dorsal side forming the

outer wing. On the outer ventraI side a short posterior

ridge tapers and ends at the posterior margin. The un

denticulated ridge is marked by a narrow groove .

On the dorsal side , anterior to the basal portion, the outer margin is concave, ending in the large, almost straight falx.

A transverse fold to the left of the basal furrow is found

in approximately half of the studied jaws. The fold is of ten

combined with a low elevated flange .

Right MIl, dorsal side : Length 0 .26--0 .68 mm, width

about half the length. A fair ly small pre-cuspidal denticle

forms the anteriormost part of the jaw and continues as a

small keel onto the cusp. The cusp is large , in cross-section

elongated, with a swollen basal part characteristically con

tinuing into the jaw, and directed almost perpendicularly

to the direction of the denticulated inner margin. AIong

the slightly convex inner margin there are no distinct inter

mediate denticles; all denticles slanting posteriorly. The

num ber of denticles varies from 9 on a 0 .26 mm longjaw to

1 3 on one that is 0 .68 mm long. These denticles are largest

in the centre of the dentition. A very narrow inner wing is

present on the posterior 3/4 of the jaw. The shank occupies

somewhat more than half the jaw length, tapering posteri

orly, and its posterior extremity is directed somewhat later-


ally towards the outer side, making the outer margin

slightly concave . The angle of the ram al arch is acute to

almost straight. The ram us is short, slightly bent upwards

and tapers to the fairly sharp tip. The anterior outer margin

forms a smooth, characteristic arch which posteriorly con

tinues into a weakly developed sinus at the base of the

ramus.


occupies the posterior 0 .6--0 .7 of the jaw length, its anterior

part surrounded by a rim which is not visible in ventrai view

but follows the edge . The opening is enclosed by almost

straigh t borders on two sides, forming an angle of 70-80° .

A high, short and narrow ridge is present at the posterior

extremity of the shank.

Left MIl, dorsal side : Length 0. 1 6--0 .64 mm, width some

what less than half the length . The cusp, with an almost

straight anterior margin, and lacking pre-cuspidal denti

cles, is strong and directed perpendicularly to the length

axis of the slightly convex, denticulated ridge . The interme

diate dentary, posterior to the cusp, is represented by 3-7

denticles of equal size. These are followed by 9-1 1 signifi

cantly larger den ticles (except for the posteriormost ones) ;

the number of denticles varies and does not seem to follow

jaw size. The posterior denticles slant; the central ones are

the largest. The inner wing is laterally extended, convex

and with its widest part at the anterior. It is more than half

the total length of the jaw in smaller specimens and less in

larger ones. The shank, occupying about half the jaw

length, tapers posteriorly, with its posteriormost tip slightly

directed towards the outer side. The outer margin , poste

rior to the ramus, is straight to slightly concave . The ram us

is fairly short, somewhat less than half of the width of the

jaw, projecting laterally, at nearly right angles and slightly

bent upwards. The anterior outer margin is smoothly sig

moidal with an almost rounded anterior part.

Ventrai side : The enclosed myocoele opening, sur

rounded by a rim, is very similar to that of the right MIl,

except for the large inner wing and the fairly low ridge in

the posterior extremity. MIll, unpaired, dorsal side: Length 0 . 1 9-0.3 1 mm, width

about half the length . The narrow, arcuate jaw has a convex

inner margin and a concave outer one. The transverse cross-section is high and fairly narrow anterior to the cusp,

and low and narrow posterior to it. The ram al angle is

about 70° . The ramal extremity is pointed and bent inwards posteriorly. The shank is slender, tapering posteriorly in

both lateral and vertical view. The shank extremity is

pointed, of ten slightly bent towards the inner side . The

anterior of the cusp, forming a border between the ram us

and the shank, is a large node on the inner margin. The

denticulation on the ridge forms the upper part of the jaw,

a cusp forms the uppermost part. The denticles slant, de

creasing in size outwards from the cusp. The cusp is pro

nounced, slanting slightly towards the posterior. Normally

there are three (2-4) pre-cuspidal denticles on the ramus.

The number of post-cuspidal denticles on the shank is

normally 5 (4-6) .

Ventrai side : The gaping myocoele on the ventrai side is

surrounded by a thickened rim.


Right MIV, dorsal side : Length 0 . 15-0.28 mm, width,

including the most basal part, about 0 .4 of the length. The

narrow, arcuate jaw has a convex inner margin and a

concave outer one . The cross-sections along the jaw are

almost triangular, high and fairly narrow. No pronounced

cusp. Denticles, slanting slightly towards the posterior, usu

ally vary in number between 6 and 8. The second and third

anteriormost denticles are usually the largest, and the size

of the following denticles decreases to the posterior end.

Left MIV, dorsal side : A mirror image of the right MIV.

Carriers: Chisel-like , flattened, almost bisymmetrical, 3-

4 times as long as the maximum width at the pronounced

head. The anterior margin of the head is straight and

abruptly truncate . The head tends to grow more asymmet

rical in later stages of ontogeny, and a shallow furrow on

one side of the head, perpendicular to the length of the

jaw, becomes more pronounced on some specimens. The

head tapers to the posterior and extends into a shaft which

in turn tapers posteriorly, ending in a pointed extremity.

The inner margin is straight. A longitudinal, shallow fur

row is present on the shaft on the opposite side of the

furrow in the head.

Remarks. - The transverse fold found on about half of the

left MI is of ten combined with a low elevated flange (Fig.

510) . This indicates that the fold may be caused by com

paction, though the overall preservation of the jaws does

not entirely support such a conclusion. A less likely expla

nation is that the fold is an expression of dimorphism.

Comparison. - The MI's show a slight resemblance to the

corresponding element of the morph with the straight

inner margin of Kettnerites ( K. ) kuberti ( see 'Comparison'

under K. (K.) kubertx) .

Kettnerites (Aeolus) microdentatus Bergman 1987 Figs. 1 21 ,J , Q, R , 1 81, 54, 55A, D-F

Synonymy. - 0 1 987 Kettnerites (Aeolus) microdentatus n. sp. -

Bergman, p. 89-9 1 , Figs. l 21 , J, Q, R, 1 8, 54, 55A, D-F.

Derivation of name. - Greek mikros and Latin dentatus, refer

ring to the minute denticles.

Holotype. - LO 5835 :2 , right MI, Fig 55E:2 .

Type locality. - Karne 3.

Type stratum. - Eke Beds, basal part.

Material. - Figs. 2 and 3; more than 250 right MI, 250 left

MI, 250 right MIl, 250 left MIl.

Occurrence. - Figs. 4 and 8; Late Llandovery to Late Ludlow,

Lower Visby Beds to Hamra Beds, unit b. Amlings l , Aurs

viken l , Bjarges l , Bodudd l , Bottarve 2, Buske l , Follingbo

2, Gannor l and 3, Gerete l , Glasskar l and 3 , Gothems

hammar 2, Haftingsklint 4, Hide l , Kalldar l , Karne 3,

Kroken 2 , Kullands l and 2 , Likmide 2 , Lilla Hallvards 4,

Narshamn 2 and 3 , Nyhamn 5, Slitebrottet 2 , Smiss l, Snack

l , Snoder 2 , Sproge 4, Stora Mafrids 2 , Stave l , Valle 2,

Valleviken l , Valve 3 , Vike 2.

Silurian paulinitid polyckaetes from Gotland 99

Diagnosis. - Right MI: Long slender jaw with about % of the

inner margin covered by fine to very fine, comb-shaped,

densely spaced denticles. Basal portion wide. Shank

pointed.

Left MI: Long slender jaw, inner margin straight, dentic

ulated along about �3 of its length. luner wing almost

rectangular, parallei to the undenticulated ridge .

Right MIl: One or two small pre-cuspidal denticles, form

ing a swelling of the cutting edge . Shank and ramus fairly

long and shank slender.

Left MIl : Fairly large cusp, long and slender shank and

ramus, inner wing narrow.

Description. - Right MI, dorsal side: Length 0.36-l .05 mm,

width about lf4-lf3 of length . The basal part is wide anterior

to the basal portion, but tapers anteriorly into a fang. The

sickle-shaped falx has a distinct cutting edge along its outer

margin. The inner margin is almost straight, covered for

about 3/4 by around 40 thin, densely spaced denticles, from

the fakal arch and posteriorly. The number, size and ex

tension of the denticles vary, from those of almost normal

size (e .g. Upper Visby Beds, Fig. 54F:2) to needle-shaped

denticles (e.g. Burgsvik Beds, Fig. 55A: l ) covering the

whole of the inner margin . The transition to the high and

narrow undenticulated ridge is fairly smooth . The ridge

ends at the posteriormost part of the shank. The inner wing

is strongly downfolded, nearly triangular, widest at its ante

rior part, with its inner margin almost straight and its

anterior corner rounded. The basal portion is fairly wide.


The basal furrow is of normal size but fairly deep. The

flange is large, thin, posteriorly highly elevated and down

folded along the posterior, outer margin, with a ligament

rim along the outer margin of the basal portion . The basal

angle is 20-30° . The shank is almost triangular with a

pointed proximity. The bight is deep, with a bight angle of

almost 90° . The outer margin , anterior to the basal portion

and posterior to the falx, has a more or less pronounced

concavity.


myocoele opening is lf4-lf3 of the jaw length. It is bounded

along the anterior and inner margins by a fairly wide and

slightly raised ligament rim. Left ML dorsal side: Length 0.38-1 . 1 8 mm, width almost

1/3 to 1/5 of length . The slender jaw tapers to the sickle

shaped falx. The fang, with a pronounced cutting edge,

follows the outer margin along the falx. The inner margin

is straight, with denticles of small to almost normal size along the anterior lf2-�3 of the inner margin between the

fakal arch and the undenticulated ridge . The anteriormost

denticles are oriented to the anterior or obliquely to the

length axis of the jaw, the posterior ones slightly to the

posterior. The denticulation decreases in size to the poste

rior, ending as small knobs. The transition onto the un

denticulated ridge is smooth. The basal portion is angular,

irregularly rectangular, with almost straight sides. The

inn er wing is almost rectangular and parallei to the un

denticulated ridge . The basal furrow is fairly deep and of

normal size, the basal angle about 20-30° . The ligament

rim is visible along the posterior outer margin of the basal

1 00 Claes F. Bergman FOSSILS AND STRATA 25 ( 1 989)

FOSSILS AND STRATA 25 ( 1 989) Silurian paulinitid polychaetes from Gotland 1 0 1

Fig. 55. Kettnerites (A.) microdentatus, except Bl-B2, K (A. ) fjaelensis, and Cl-C2, K (A.) cf. fjaelensis. All specimens are i n dorsal view, x120 except Al , A2, E3 and E4 . DA. Narshamn 2, Burgsvik Beds, 83-l 2LJ; Al left MI , LO 5831 : 1 , x80 ; A2 right MI , LO 5831 :2, x80. DB. Kullands 2, Hemse Beds, Hemse Mari , NW part, 84-3l 2DF; Bl left MI, LO 5832:5 ; B2 right Ml, LO 5832:6 . D C. Viclfålle l, Hemse Beds, unit b, 75-41 CB; C l left MI , LO 5825: 1 ; C2 right MI , LO 5825:2 . OD. Gannor l , Eke Beds, basal part, 7l-l 23LJ; Ol left MI, LO 581 1 :3 ; D2 right MI , LO 581 1 :4 . D E. Kårne 3, Eke Beds, basal part, 7 l-199LJ; El left MI, LO 5835 : 1 ; E2, holotype, right MI, LO 5835:2 ; E3 1eft MIl, LO 5835:3 , x2 1 O; E4 right MIl, LO 5835:4, x2l0. O F. Gannor 3, Hemse Beds, Hemse Mari, SE part, 7l -l25LJ; Fl left MI, LO 581 3: 1 ; F2 right MI, LO 5813 :2 .

Fig. 54 (opposite side) . Kettnerites (A. ) microdentatus. All specimens are in dorsal view, x1 20. DA. Vattenfallsprofilen l , Hogklint Beds, unit b , 70-6LJ; Al left MI, LO 5766: 1 ; A2 right MI , LO 5766:2 ; A3 left MIl , LO 5766:3; A4 right MIl, LO 5766:4. DB. Lickershamn 2, Lower Visby

Beds, unit f, 73-53LJ; Bl left MI, LO 5763:2; B2 right MI, LO 5763:3 . DC. Vike 2, Slite Beds, Pentamerus gothlandicus Beds or slightly older, 83-4CB; Cl left MI, LO 578 1 : 1 ; C2 right MI, LO 578 1 :2 . O D. Stave l, Slite Beds, Slite Mari, central part, 75-1 1CB; Ol left MI, LO 5772: 1 ; D2

right MI, LO 5772 :2; D3 1eft MIl, LO 5772 :3; D4 right MIl, LO 5772:4. D E. Vattenfallsprofilen l, Hogklint Beds, unit b, 70-6LJ; E l left MI, LO 5766:5 ; E2 right MI, LO 5766:6; E3 right MIl, LO 5766:7. OF. Håftingsklint l, Upper Visby Beds, 76-9CB; Fl left Ml, LO 5367: 1 ; F2 right MI, LO 5367:3; F3 1eft MIl, LO 5367:2; F4 right MIl, LO 5367:4. O G. Vattenfallsprofilen l, Hogklint Beds, unit a, 70-20LJ; Gl left MI, LO 5764:4; G2 right MI, LO 5764:5. OH. Vattenfallsprofilen l, Hogklint Beds, unit d, Valdaria testudo leve!, sample RM; Hl left MI, AN 2717; H2 right MI, AN 271 8.

1 02 Claes F. Bergman

portion. Anterior to the basal portion there is a wide and

very shallow concavity on the outer margin .

VentraI side : The crescent-shaped, strongly enclosed

myocoele opening is about lf4-lf3 of the jaw length and

resembles the one of the right MI.

Right MIl, dorsal side: Length 0 . 1 7-0.68 mm, width nor

mally more than half the length. There are two (in some

populations probably only one) pre-cuspidal denticles on

the dorsal side of the cusp, forming a swelling of the cutting

edge, which does not reach the anteriormost part of the

jaw. The anterior margin is sickle-shaped, forrned by the

large cusp which slants towards the posterior. The succeed

ing denticle is small; the transition to the almost straight

denticulated ridge is gradual. Normally there are 6--8 slant

ing denticles on the ridge ; these decrease slightly in size to

the posterior. The shank is long and slender, occupying

more than half the length of the jaw, and tapering posteri

orly. !ts extremity is almost pointed and bent slightly ven

trally outward. The outer margin of the shank is almost

straight. The bight is fairly deep, the bight angle acute to

almost straight. The ramus is fairly large and slender, its

extremity rounded. The anterior outer margin is almost

straight.


�3-3j4 of the jaw length. The almost straight anterior and

inner margins are surrounded by a ligament rim of normal

size. The opening extends anteriorly in the anterior inner

part, forming a crescentic shape.

Left MIl, dorsal side: Length 0 .31-0.53 mm, width ap

proximately half the length . The large cusp forms the

sickle-shaped anterior margin. The intermediate dentary is

distinct, with 4-7 denticles slightly increasing in size to the

posterior. The denticles, normally 6--8, slant to the poste

rior on the slightly convex, denticulated ridge . The denti

cles increase in size to a maximum at the middle of the

dentary whence they dec re ase to the posterior, ending

fairly abruptly (e.g. in specimens from Snoder 2) or more

gradually (e.g. in those from Håftingsklint l ) . The shank

occupies about half the length of the jaw, tapers posteriorly

and ends fairly sharply. The inner wing is very narrow and usually not visible in dorsal view. The outer margin of the

the shank is straight to slightly concave . The bight is of

moderate size, the bight angle almost straight to slightly acute . The anterior outer margin is smooth and vaguely

sigmoidal .


about lf2�3 of the jaw length . The opening is surrounded

by a flat ligament rim. The inner and anterior margins are

alm ost obliquely oriented to one another.

Remarks. - The basal portion of the right MI of the popula

tion of Narshamn 2 is conspicuously wide with a prominent

flange, but whether this is the main trend in the lineage or

just an atypical form developed in a population, can at

present not be deduced because of the shortage of mate

rial .

There are 1-2 pre-cuspidal denticles of the right MIl.

Normally the dentition formula of the anteriormost denti

cles of the MIl evolves very slowly in a lineage . We may here

have an exception, but it is also possible that two different


lineages occur in paralleI. The difference between them is

very modest and may be summarized as follows: In the

robust form the left and right MI are somewhat shorter with

slightly larger denticles; the right MIl has one pre-cuspidal

denticle and gives a more short and wide impression than

the MIl of the slender type.

Comparison. - The left and right MIl of K. (A. ) microdentatus

are very similar to the corresponding element of K. (A. )

sisyphi sisyphi and K. (A. ) fjaelensis. The Ml ' s however, are

diagnostic.

Kettnerites (Aeolus) siaelsoeensis Bergman 1987 Figs. 1 2H, 1 8K, 56

Synonymy. - 0 1 987 Kettnerites (Aeolus) siaelsoeensis n. sp . -

Bergman, p. 91-93, Fig. 1 2H, 18 , 56.

Derivation of name. - Latin Siaelsoeensis, from Sjalso, referring

to the type locality.

Holotype. - LO 5360:2 , right MI, Fig 56B.

Type locality. - Sjalso l . Type stratum. - Lower Visby Beds, unit b.

Material. - Fig. 2 ; more than 80 right MI, 80 left MI, 62 right

MIl, and 49 left MIl.

Occurrence. - Figs. 4 and 7; Late LIandovery. Lower Visby

Beds, unit b. Gustavsvik l , 2, and 3, Ireviken 3, Ronnklint l , Sjalso l . Diagnosis. - Right MI: Slender, tapering towards the

pointed fang; straight inner margin with thin comb-shaped

dentary; shank sharp-ended. Basal plate almost triangular,

fused to jaw.

Left MI: Slender, tapering towards anterior end, comb

shaped dentary ending before reaching the low and narrow

undenticulated ridge . Basal furrow small and deep, extending somewhat under the undenticulated ridge .

Right MIl: Large , sickle-shaped cusp with a fairly small

pre-cusp on its anteriormost part. Very slender shank which tapers to the posterior.

Left MIl: Large slender cusp, shank slender, tapering to the posterior, diminutive inner wing.

Description. - Right MI, dorsal side: Length 0 .34-0.62 mm,

width about or slightly less than 1f4 of length. The slender

jaw tapers to a thin, sickle-shaped and fairly long fang,

slightly bent upwards in relation to the almost flat jaw

surface. The inner margin is straight and covered by thin,

long, perpendicular denticles (26 on a 0.34 mm long jaw;

42 on a 0 .62 mm long one) from the fakal arch to the

undenticulated ridge . The latter is straight, high and nar

row, almost forming the posteriormost sharp tip of the

shank. The inner wing is alm ost rectangular, strongly down

folded, with an almost straight inner margin. The basal

furrow is partly covered by the almost triangular, undentic

ulated basal plate . The inn er margin of the basal plate is

smoothly rounded; the posterior, outer part is occupied by


a fold. The flange is rounded, the ligament rim exposed as

a very indistinct, narrow furrow, to the right of which is a

low fold. The basal portion is almost rectangular; the basal

angle approximately 20° . Anterior to the basal portion the

outer margin is smoothly concave along the posterior half

of the jaw, whence it swings out forming the sickle-shaped

falx.

Ventrai side : The strongly enclosed, rounded myocoele

opening is about 1/3 of the jaw length . Along the anterior,

outer margin of the basal portion the ligament rim is a

smoothly rounded ridge . Along the inner wing the ridge is

narrow and surrounded by a groove .

Left MI, dorsal side : Length 0.32-0.61 mm, width lf4 of

length . The slender jaw tapers to a long fang which is

slightly bent upwards in relation to dorsal surface . The

inner margin is straight, with 20-50 needle-like denticles in

the comb-shaped dentary. The denticulation covers most

of the inner margin behind the falcal arch, ending as

crenulations before reaching the inner wing. There is a

smooth transition to the undenticulated ridge which is

straight, low and narrow, ending close to the almost

straight posterior margin. The inner wing is of normal size,

0 .20-0.25 of the jaw length, almost rectangular and moder

ately downfolded. The basal furrow is short and deep,

projecting under the undenticulated ridge . The outer mar

gin of the basal portion is smoothly rounded, slightly down

folded, extending for a quarter of the jaw length, with the

iigament rim forming a flat fold posteriorly. The basal

portion is almost rectangular and smoothly rounded, the

basal angle is about 20° . The outer margin anterior to the

basal portion has a fairly deep concavity at the middle of

the jaw.


almost a mirror image of that in the right MI.


about half the length . A fairly small pre-cuspidal denticle ,

on the cutting edge of the anterior margin of the sickle

shaped cusp, forms the anteriormost part of the jaw. The

cusp points towards the posterior. It is followed by two

intermediate denticles, the anteriormost of which is

smaller, and about 8-1 0 denticles of moderate size which

decreases towards the posterior. The shank is long, more

than half the jaw length, and very slender. The inner wing

is diminutive . The inner and outer margins of the shank are convex and concave, respectively, the posteriormost

part ends fairly sharp ly, pointing slightly laterally outwards.

The bight is deep, the bight angle acute . The ramus is fairly

long and somewhat needle-like , with only a trace of a sinus

on the anterior, outer margin .


�3-3f4 of the jaw length . The opening extends anteriorly, i s

delimited to the anterior and inner side by the almost

straight margins forming a sharp angle. The ligament rim

is narrow and only slightly widened on the outer part of the

ramus.

Left MIl, dorsal side : Length 0.29-0.49 mm, width ap

proximately half the length . The large, fairly slender,

sickle-shaped cusp is followed by 6-7 denticles of different

sizes and orientations. The anteriormost ones point anteri-


Fig. 56. Kettnerites (A . ) siaelsoeensis. All specimens from Sjalso l , Lower Visby Beds, unit b, 79-1 2L]. All speeimens are in dorsal view, x1 20. DA. Left MI, LO 5360: 1 . DB . Holotype, right MI, LO 5360:2 . De. Left MIl , LO 5360:3 . OD. Right MIl , LO 5360:4. DE. MIll (unpaired) , LO 5360:5 . OF. Right MIV, LO 5360 :6. OG. Left Ml, LO 5360:7. OH. Left Ml!, LO 5360:8.

orly, the ones be hind are posteriorly oriented. The follow

ing 7-8 denticles are of moderate size, slanting, and decrease in size towards the posterior. The inner wing is very

narrow, about half as long as the jaw. The slender shank,

about half the jaw length or slightly less, tapers posteriorly

to a fairly sharp end. The bight angle is usually acute , in

some cases straight. The ram us is fairly long and slender,

with a pronounced sinus at the anterior outer margin .

Ventrai side: The crescent-shaped, slightly enclosed mya

coele opening is somewhat more than half the jaw length .

The opening extends slightly to the anterior inner side of

the jaw. The almost straight margins are surrounded by a

ligament rim of normal size.


Remarks. - The good preservation of the type collection,

probably due to low water energy and low bioturbation

leveis, made it possible to make a fairly safe identification

of MIll and MIV. These jaws are more or less identical to

those of K. (A.) sisyphus klasaardensis (see description of this

species) .

Comparison. - The form and den ticulation of the MI are very

similar to the corresponding element of K. (A. ) micro

dentatus, except for the fused basal plate . The MIl of K. (A. )

siaelsoeensis is usually more slender and has a single pre-cusp

on the right MIl, while there is usually a double pre-cusp

on the right MIl element of K. (A. ) microdentatus.

Kettnerites (Aeolus) fjaelensis Bergman 1987 Figs. ] 2P, ] 8J, 55B, c, 57

Synonymy. - O 1 987 Kettnerites (Aeolus) fjaelensis n. sp . - Berg

man, p. 93-95, Figs. 1 2P, 1 8, 55B, C, 57.

Derivation of name. - Latin fjaelensis, referring to the type

locality Fjåle 3 .

Holotype. - LO 5798:2 , right MI, 57D:2 .

Type locality. - Fjåle 3 .

Type stratum. - Klinteberg Beds, unit e , close to the Wen

lock-Ludlow boundary.

Material. - Fig. 3; more than 28 right MI, 30 left MI, 19 right

MIl, 1 6 1eft MIl.

Occurrence. - Figs. 4 and 8; Late middle Wenlock to Early

middle Ludlow, Halla-Eke Beds. Fjåle 3, Gandarve 1 , Herr

vik 2, Hide 1 , Horsne 5, Kullands 2, Likmide 2, Lukse 1 ,

Mulde Tegelbruk 1 , Snoder 2 , Tomtbodarne 1 , Vidfålle 1 .

Diagnosis. - Right MI: Slender jaw, tapering strongly to the

large fang. The anterior two thirds of the inner margin

covered by fairly wide-spaced, anteriorly fairly large , almost

triangular denticles. Basal portion wide with conspicuously large, rounded flange. Shank pointed.

Left MI: Mirror image of the right MI with the exception of the basal portion (no bight or basal plate) . Inner wing

almost rectangular.

Right MIl: Two very small to fairly small pre-cuspidal

denticles. Cusp moderately large , slightly swollen at its base .

Left MIl : Single , fairly large cusp. Shank and ramus long

and slender.

Description. - Right MI, dorsal side: Length 0 .33-0.76

mm, width slightly more than half the length . The jaw is

very wide at its base anterior to the basal portion and tapers

anteriorly, ending in the fang. The large sickle-shaped falx

has a distinct cutting edge along its outer margin. The

inner margin is almost straight and covered by about 1 5-20

slightly spaced, triangular denticles over about �3 of the

inner margin from the falcal arch. The anteriormost 5-1 0

denticles are of normal size, but the size decreases to the

posterior, where the denticles are ultimately reduced to

small knobs. There is a smooth transition to the high and

narrow undenticulated ridge which ends at the posterior

most part of the shank. The inner wing is very deeply


downfolded, almost triangular, widest at the anterior, its

inner margin almost straight and its anterior corner

rounded. The basal portion is wide, its outer margin

rounded.


The basal furrow is of normal size but fairly deep. The

flange is very wide , large , thin, posteriorly highly elevated

and slightly downfolded along the rounded, posterior,

outer margin. The ligament rim along the outer anterior

margin of the basal portion is more or less indistinct. The

basal angle is about 20-35° . The shank is slender and

almost triangular with a pointed extremity, slightly bent to

the outside in relation to the denticulated ridge . The bight

angle is almost 90° . The transition between the anterior

most part of the basal portion and the outer margin is

rounded. A minor concavity is present posterior to the falx.


myocoele opening is about 1/3 of the jaw length, or slightly

less. The ligament rim is fairly distinct and forms a low,

sharp ridge on the inner wing.

Left MI, dorsal side : Length 0 .42-0.67 mm, width about

1/4 of length . The slender jaw tapers to the sickle-shaped

falx. The fang is large , with a pronounced cutting edge

along the outer margin of the falx. The inner margin is

straight, with slightly spaced denticles ofnormal size on the

anterior of the inner margin. The denticles cover slightly

more than half of the inner margin between the falcal arch

and the undenticulated ridge . They decrease in size poste

riorly, ending as small knobs. The anteriormost denticles

project anteriorly or perpendicularly to the length axis of

the jaw, the posterior ones are more or less perpendicular

or point somewhat posteriorly. The transition to the un

denticulated ridge is very smooth . The basal portion is

wide, irregularly rectangular, with almost straight inner

margin, slightly concave posterior side and a slightly convex

to rounded outer side . The inner wing is almost rectangu

lar, paralleI to the undenticulated ridge . The basal furrow

is of normal size, the basal angle about 25-40° . The liga

ment rim along the posterior, outer margin of the basal

portion is indistinct. In some populations an outer wing is developed (Fig. 57D: 1 , B: 1 ) . The outer margin anterior to

the basal portion is almost straight, ending in the falx.


myocoele opening is about 1/3 of the jaw length or slightly less. The opening is similar to that of the right MI.

Fig. 57. Kettnmtes (A.) fjaelensis. All specimens in dorsal view, x] 20, except B4. DA. Mulde Tegelbruk l, Mulde Beds, 82-7CB; Al left MI, LO 5788:2 ; A2 right MI, LO 5788: l ; A3 Ieft MIl, LO 5788:3; A4

right MIl, LO 5788:4. DB. Snoder 2, Hemse Beds, Hemse Marl NW part, 82-I 4CB; Bl left MI, LO 5809:3; B2 right MI, LO 5809:4; B3 left MIl, LO 5809:5; B4 right MIl, LO 5809:6, x lOO. DC. Horsne 5, Halla Beds, unit b, 75-52CB; C I left MI, LO 5794: 1 ; C2 right MI, LO 5794:2; C3 left MIl, LO 5794:3; C4 right MIl, LO 5794:4. OD. Fjale 3, Klinteberg Beds, unit e , 77-7CB; Dl left MI , LO 5798 : 1 ; D2 , holotype, right MI , LO 5798:2 ; D3 left MIl, LO 5798:3; D4 right MIl, LO 5798:4. DE. Gandarve 1, Halla Beds, 7 l-8l CB; E I left MI, LO 5836:2 ; E2 right MI, LO 5836:3. OF. Likmide 2, Hemse Beds, Hemse Marl, SE part, 82-28LJ; Fl left MI, LO 580 1 : 1 ; F2 right MI, LO 580] : 2 .



Right MIl, dorsal side: Length 0.30-0 .47 mm, width

about half the length or slightly more . Two pre-cuspidal

denticles form the anteriormost part of the jaw. Their size

varies from very small, indistinct swellings of the cutting

edge (e.g. Fig 57C:4) to fairly large, separate pre-cusps (Fig.

57D:4) . The anterior margin is more or less rounded,

bearing a large cusp with a swollen base . The cusp is di

rected perpendicular to the length axis of the jaw or slants

slightly to the posterior. The following 1-2 denticles are

small . There is a gradual transition to the almost straight

denticulated ridge, with a post-cuspidal dentary composed

of 8-1 0 slanting denticles, largest at the centre . The shank,

about half the length of the jaw, is slender and tapers to the

posterior; its extremity is pointed and bent slightly ventrally

outward. The bight is of normal size, the bight angle almost

90' . The ramus is large , wide, almost triangular and some

what pointed. The anterior outer margin is almost straight.

VentraI side : The slightly enclosed myocoele opening is

1j2-�3 of the jaw length. The ligament rim is wide with a low

relief.

Left MIl, dorsal side: Length 0.32-0 .47 mm, width ap

proximately half the length. The fairly large cusp with a

distinct cutting edge forms the sickle-shaped anterior mar

gin . The intermediate dentary is distinct with 3-5 fairly

small denticles of more or less equal size. The usually 8-1 0

post-cuspidal denticles slant to the posterior, the largest at

the middle. The shank, about half (or slightly less) of the

jaw length, tapers to a fairly sharp posterior end. The inner

wing is very narrow and usually not visible in dorsal view.

The outer margin of the shank is straight to slightly con

cave. The bight is of moderate size, the bight angle almost

straight to slightly acute . The ramus is almost triangular,

wide at its base . The anterior, outer margin has a slight

concavity at the centre .

VentraI side : The crescent-shaped, slightly enclosed myo

coele opening is about % of the jaw length. The opening

extends to the anterior on the anterior, inner side of the

jaw.

Comparison. - The MI ' s are easy to identify on the denticu

lation . The MII ' s are very similar to the corresponding element of K. (A. ) sisyphi sisyphi and K. (A. ) microdentatus.

Genus Lanceolatites Bergman 1987

Synonymy. - Lanceolatites n. gen. - Bergman, p . 96 .

Derivation ofname. - Latin lanceolatites, from lance, referring

to the very elongated and sharp-pointed MI.

Type speeies. - Lanceolatites gracilis Bergman 1 987

Other species. - L. sp. A.

Diagnosis. - Right MI: Long slender jaw, oval cross-section,

comb-shaped dentary, pointed shank.

Left MI: Long slender jaw, oval cross-section, comb

shaped dentary, small basal portion, almost rectangular

inner wing.

Right MIl: Double cusp with one minor denticle between

the cusps; shank and ramus long, slender with pointed

extremities. Bight deep.


Left MIl: Double cusp, shank and ramus long, slender

with pointed extremities.

Remarks. - Lanceolatites is characterized by the conspicu

ously long, slender and pointed extremities of its jaws, the

slenderness of ten being combined with an undulating

shape, particularly the late Wenlock to Ludlow Lanceolatites

gracilis specimens. The type species has a variety which is

coarser and of ten slightly smaller than the type form. The

Lanceolatites sp. A specimens are less slender and have fewer

denticles.

Lanceolatites gracilis Bergman 1 987 Figs. I4A, B, I 8L, I 9C, 58, 5 9 , 60B-D

Synonymy. - D 1 979 'Oenonites ' aspersus Hinde - Bergman, p.

99, Fig. 28:4C,D (only) . D 1 987 Lanceolatites gracilis n. sp. -

Bergman, p. 96-100, Figs. 1 4A, B, 1 8, 1 9C, 58, 59, 60B, C,

D .

Derivation of name. - Latin gracilis, referring to the long

slender form of both MI and MIl .

Holotype. - LO 5365:4, right MI, Fig. 58D :2 .

Type locality. - Nygårdsbackprofilen l . Type stratum. - Lower Visby Beds, unit e .

Material. - Figs. 2 and 3; more than 171 right MI, 1 58 left

MI, 1 76 right MIl, 1 2 1 left MIl.

Occurrence. - Figs. 4 and I l ; Late Llandovery to latest midd le

Ludlow, Lower Visby Beds, unit e , to Eke Beds, lower-mid

dIe part. Ajmunde 1 , AmIings 1 , Autsarve l , Bodudd l , 2 ,

and 3 , Buske l , Fågelhammar 3, Fjale 3 , Gannor l and 3,

Gardsby l , Gerete 1 , Gerumskanalen l , Gnisvard l and 2 ,

Gogs 1 , Grogarnshuvud l , Haftingsklint 1 and 4, Hagvide

1 , Hummelbosholm 1 , Ireviken 3, Kalldar l , Kattelviken 5 ,

KJasård l , Korpklint 1 , Kullands 2 , Lickers 1 , Lickershamn

2, Likmide 2, Lilla Hallvards l and 4, Lukse l , Nar 2 ,

Nygårdsbackprofilen l , Petsarve 2 , Ronnklint l , Snackgardsbaden l , Snoder 2 , Stora Kruse l , Vaktård 2 , 4 , and 5 , Vastlaus l , Vattenfallsprofilen 1 .

Diagnosis. - Right MI: Long, slender, evenly tapering jaw,

almost straight inner margin, covered along its entire

!ength by about 50-60 denticles in a comb-shaped dentary.

Pointed shank.

Left MI: Apart from the small, angular basal portion with

a nearly rectangular inner wing, it is almost a mirror image

of right MI.

Right MIl : Double cusp with two parts of almost equal

size, separated by a minor denticle. Three to six intermedi

ate denticles of equal size. Shank and ramus long, slender,

with pointed extremities.

Left MIl: Double cusp, parts of equal size. Intermediate

denticles covering up to half the length of the denticulated

inner margin. Shank and ramus long and slender with

pointed extremities.

Description. - Right MI, dorsal side: Length 0 .36--1 . 1 8 mm,

width lfs of length. The slender jaw tapers anteriorly, end

ing in a narrow, pointed fang, which is bent fairly strongly


Fig. 58. Lanceolatites gracilis. E and F L. gracilis var. visby. All specimens in dorsal view; all x80 except C. DA. Left M I , LO 5765:2 , Lickershamn 2, Hogklint Beds, unit a, 73-57Lj. DB . Right MI, LO 5764:3, Vattenfallsprofilen l, Hogklint Beds, unit a, 70-20Lj. DC. Same sample as A; Cl

left Mil, LO 5765:4, x1 20; C2 right Mil , LO 5765 :5, x120 ; C3 left Mil , LO 5765:6. OD. Nygårdsbåckprofilen l , Lower Visby Beds, unit e , 79-42LJ; D l left MI, LO 5365:3; D2, holotype, right MI, LO 5365 :4; D3 1eft Mil , LO 5365:5; D4 right MIl, LO 5365 :6. DE. Vattenfallsprofilen l, Lower Visby Beds, unit e ( ? ) , 76-6LJ; El left MI, LO 5366 : 1 ; E2 right MI, LO 5366:2; E3 left Mil, LO 5366:3; E4 right Mil, LO 5366:4. OF. Vattenfallsprofilen l , Hogklint Beds, unit b , 70-6LJ; Fl left MI, LO 5 768: 1 ; F2 right MI, LO 5768 :2; F3 left MIl , LO 5768: 3; F4 right Mn, LO 5768:4.

1 08 Claes F. Bergman FOSSILS AND STRATA 25 ( 1 989)

Fig. 59. Lanceolatites gracilis, Hemse Beds, Hemse Mari, NW part, all speeimens in dorsal view, x 1 20. DA. Likmide 2, 82-28LJ; Al left MI, LO 5802: 1 ; A2 right MI, LO 5802:2 ; A3 left MIl, LO 5802:3 ; A4 right MIl, LO 5802:4. DB. Ajmunde l , 7 1-40LJ; B l left MI, LO 5807: 1 ; B2 right

MI, LO 5807: 2; B3 left MIl, LO 5807:3; B4 right MIl, LO 5807:4.

Fig. 60 (opposite page) . All speeimens in dorsal view, except EI-4, all x1 20, except A, D2, D3, E l and E2. DA. Lanceolatites sp. A, Strands l , Hamra Beds, unit b , 75-1 4LJ; Al left MI, LO 5840:5, x80; A2 right MI, L O 5840:6, x80. O B . L. gracilis var. visby, Fjåle 3 , Klinteberg Beds, unit

e , 77-7CB; Bl left MI, LO 5797 : 1 ; B2 right MI, LO 5797:2; B3 1eft MIl, LO 5797:3; B4 right MIl, LO 5797:4. D e. L. gracilis Vaktård 4, Hemse Beds, Hemse Mari, SE part, 8 1 -35LJ; C l left Ml, LO 5821 : 1 ; C2 right MI, LO 582 1 :2 ; C3 1eft MIl, LO 582 1 :3; C4 right MIl, LO 582 1 :4 . OD. L. gracilis Kålldar l , Hemse Beds, Hemse Mari, NW part, 75-75CB; Dl right MI, LO 5810 :2 ; D2 basal portion, with basal plate in place , of right

MI, same specimen as D l , x600; D3 anterior part of left MI, LO 58 10 : 1 , x600. D L. gracilis, Våstlaus l, Hemse Beds, Hemse Mari, SE part, ventrai view, 82-3ILJ; E l , right MI, LO 5806: 1 , x21 0; E2, left MI, LO 5806:2, x2 10 ; E3, right MIl, LO 5806:3; E4, left MIl, LO 5806:4.

FOSSILS AND STRA TA 25 ( 1989) Silurian paulinitid polychaetes from Gotland 109


upwards in relation to the convex dorsal surface . The dis

tinct cutting-edge runs transversely to the extension of the

jaw along the fang. In the Late Llandovery and Early Wen

lock specimens the long (0 .7 of the jaw length) , almost

straight inner margin has about 50-60 thin , needle-shaped

denticles of equal size along its entire length (Fig. 58D:2) .

In Late Wenlock and Early Ludlow specimens the num ber

of denticles is smaller, about 50 or even lower (Fig. 58B:2 ) .

The denticulation and the inner margin end before reach

ing the inner wing. There is an abrupt transition to the

high, smoothly convex and very narrow, undenticulated

ridge . The posterior part of the ridge is somewhat broad

ened, slightly bent outward at the middle , extending down

and forming the posteriormost part of the shank together

with the inner wing. The inner wing is usually downbent,

the anterior outer corner is rounded, the inner margin

medially concave. The shank is wide posteriorly, fairly

blunt-ended, though the undenticulated ridge forms a con

spicuously pointed tip. The basal furrow is long, wide and

fairly deep. The posterior part of the elevated flange is fairly

thin-walled. There is a short and fairly narrow fold in the

outer, anterior margin of the basal portion . The basal angle

is 30-40° .

The basal plate which i s fused to the jaw, i s recorded only

from specimens of late Wenlock and Ludlow age (Fig.

60D:2 , C :2) . It is almost triangular, with a semicircular,

rounded margin along the inner side .

The outer margin, anterior to the fairly small basal por

tion is smoothly wavy with a distinct median concavity. The

fal x is sickle-shaped, very short and indistinct.

VentraI side : The strongly enclosed myocoele opening,

about 1;3 of the jaw length, is surrounded by a narrow, high

and fairly sharp ridge, the ligament rim. The opening is

anteriorly crescent-shaped, laterally following the inner

and outer margins (Fig. 60E: l ) .

Left MI, dorsal side: Length 0 .40-1 . 1 8 mm, width 1;5 of

length . The jaw tapers evenly towards the anterior, ending

in a finely pointed fang, only slightly bent upwards in

relation to the convex, dorsal surface . The fang has a

distinct, almost transverse cutting-edge. In the Late LIandovery and Early Wenlock specimens, the inner margin is

almost straight, densely denticulated by around 50-60 nee

dle-shaped denticles of equal size, oriented perpendicular

to the jaw axis. In Early Ludlow specimens the num ber of

denticles is about 50 or less. The comb-shaped dentary, about 0 .7 of the jaw length, completely covers the inner

margin and does not reach the inner wing. The transition

to the undenticulated ridge is abrupt. The ridge is slightly

elevated and rounded in longitudinal view, fairly narrow,

thinning posteriorly. The posteriormost part of the ridge is

bent inwards along the posterior margin of the deeply

downfolded inner wing. The basal furrow is of normal

!ength, wide and fairly deep. The posterior half of outer

margin of the basal portion is slightly downfolded. The

anterior half of the ligament rim is visible. The basal por

tion is almost triangular with a very small outer face . The

basal angle is about 40-50° . The outer margin , anterior to

the basal portion, is smooth, rounded in the anterior-pos-


terior direction, normally with a concavity at middle. The

falx is short and sickle-shaped.

VentraI side : The strongly enclosed myocoele opening is

about %-1;3 of the jaw length . The shape of the opening is

very similar to that of the right MI (Fig. 60E :2 ) .

Right MIl, dorsal side : Length 0 .31-0.93 mm, width

somewhat less than half the length . The almost equal parts

of the double cusp are separated by a min or denticle. The

posteriormost part slants slightly more to the posterior. In

the Late Llandovery and Early Wenlock specimens, the

intermediate dentary is well developed with about three

equal denticles of normal size; whereas specimens from

latest Wenlock to Ludlow Beds have more denticles, about

5-6. The post-cuspidal dentary is represented by 1 0-1 2

fairly slender, slightly slanting denticles. The shank tapers

to the posterior, the posterior 0 .3 be ing without denticles,

the extremity very pointed and of ten characteristically bent

to the right. The shank occupies slightly more than half the

jaw length. The bight is deep with an acute bight angle. The

ramus is long, of ten slender and pointed. The anterior

outer margin has a wide and shallow sinus.

VentraI side : The parabolic, slightly enclosed myocoele

opening extends anteriorly on the inner side for about �3

of the jaw length . The ligament rim is narrow (Fig. 60E:3) .

Left MIl, dorsal side : Length 0 .47-1 .40 mm, width less

than half the length . The double cusp has parts of equal

and moderate size. The posterior part slants slightly to the

posterior, and the anterior one is perpendicular to the

length axis of the jaw. The intermediate dentary is repre

sented by about 1 2-15 small denticles. There are about ten

post-cuspidal , fairly small, slightly slanting denticles de

creasing in size to the posterior. The denticulation ends

abruptly at the posterior end of the inner wing, !eaving the

posterior 0.2 of the shank without denticles . The fairly

slender shank, slightly less than half the jaw length, tapers

distinctly to the posterior, its posterior proximity sharply

pointed. The bight is fairly deep, the bight angle acute . The

ramus is usually fairly slender and long with a pointed

proximity. The anterior, outer margin has a wide, shallQw

sinus. VentraI side : The slightly enclosed myocoele opening,

somewhat more than half the jaw length, is slightly curved

and surrounded by a fairly narrow rim along the anterior

and the inner side (Fig. 60E:4) .

Remarks. - The transition from the intermediate to the

post-cuspidal dentary of the MIl is gradual in some popula

tions, making the count of intermediate denticles some

what uncertain .

Lanceolatites gracilis var. visby Figs . 1 4C, D, 1 8L, S8E, F, 608, D

Occurrence. - Early Wenlock to Wenlock-Ludlow boundary,

Lower Visby Beds, unit e, to Hemse Beds, Hemse Mari NW

part. Ajmunde l , Buske l , Fjiile 3, Gnisviird l , Hiiftingsklint

4, Ireviken 3 , Korpklint l , Lickers l , Nygårdsbiickprofilen

l , Nyhamn 4, Ronnklint l , Sniickgiirdsbaden l , Vattenfalls

profilen l .


Diagnosis. - Right MI: Fairly long, slender jaw, tapering

evenly to the anterior. Densely denticulated, straight inn er

margin with around 40 denticles.

Left MI: Fairly long, slender jaw, tapering evenly to the

anterior. Almost straight inner margin densely denticu

lated with around 40 needle-shaped denticles.

Right MIl: Double cusp, parts of equal size, separated by

a denticle. Two, more rare ly three, intermediate denticles.

Shank fairly slender.

Left MIl: Double cusp, parts of almost equal size, fol

lowed by 6--8 intermediate denticles. Shank fairly sJender.

Description. - Right MI, dorsal side : Length 0 .42-0.95 mm,

width about 1/5 of length . The inner margin is straight,

strongly denticulated with approximately 40 densely

spaced, erect, needJe-shaped denticles, slightly decreasing

in size posteriorly. In elements from the Lower Visby Beds,

the denticles cover the complete inner margin, 0 .6--0 .7 of

the jaw length (Fig. 58E:2 ) . In those from the Klinteberg

Beds the denticulation covers about 0 .6 of the jaw length,

ending before reaching the inner wing (Fig. 60B:2) .

VentraI side : Similar to the nominal species.

Left MI, dorsal side : Length 0.49-1 .01 mm, width about

lf4-lf5 of length . The num ber of denticles is about 40 (Fig.

60B: l ) . Except for the basal part, this is alm ost a mirror

image of the right MI.


Right MIl, dorsal side : Length 0.42-0 .70 mm, width less

than half the length. The double cusp has parts of alm ost

equal size and separated by a denticle. The anterior part,

bent upwards in relation to the dorsal surface, is perpendic

ular to the length axis of the jaw; the posterior part slants

slightly to the posterior. Two denticles of fairly small and

equal size form the intermediate dentary, sometimes with

a slightly larger denticle forming the transition to the

post-cuspidal dentary which has 7-1 0 fairly large, posteri

orly slanting denticles (Fig. 60B:4) . The shank is sJender

and tapers to the posterior, with a pointed extremity, some

times slightly bent outwards. The shank occupies about half

the jaw length. The bight is fairly deep, the bight angle

acute ; the ramus is almost triangular. The anterior outer

margin is almost straight, ending in a pointed anterior end. VentraI side : Similar to the nominal species.

Left MIl, dorsal side : Length 0.40-0 .75 mm, width less

than half the jaw length . The double cusp has parts of

almost equal size and varying orientation: usually the ante

rior part is oriented slightly anteriorly (Fig. 58E:3) or per

pendicularly to the length axis of the jaw, and the posterior

one slightly posteriorly or perpendicularly to the length

axis. There are 6--9 intermediate denticles and a somewhat

gradual transition to the post-cuspidal dentary with 6--1 1

posteriorly slanting denticles. The shank, slightly less than

half the jaw length, tapers to a pointed posterior end. The

inner wing is narrow. The bight is fairly deep, the bight

angle acute . The ramus is almost triangular, with a some

what pointed extremity. The anterior outer margin is al

most straight or has a very weak, wide sinus. The anterior

end is alm ost sickle-shaped.


Silunan paulinitid polychaetes from Gotland I I I

Discussion. - The MI 's are very similar to those of L. gracilis,

but the latter are slightly more sJender and have a larger

number of denticles. The MIl ' s differ more, both in gen

eral morphology (L. gracilis being more sJender) and in the

number of denticles.

The species evolved slowly; changes of denticle formulae

of the MIl element are fairly easy to observe . There is no

verifiable size difference between them, such as would be

expected if they had been ontogenetic variations of the

same species. Also, they are only occasionally found to

gether. L. gracilis has a longer stratigraphical range than

the variety, so they are not likely to be sexual dimorphs.

The variety seems always to be less frequent than the nom

inal species.

Lanceolatites Sp . A Figs. 1 4E, 60A

Material. - Nine right MI, 1 2 1eft MI, 1 3 right MIl, 8 left MIl.

Occurrence. - Fig. 4 and I l ; Late Ludlow, Hamra Beds, unit

b. Bankvat l, Strands l .

Diagnosis. - Right MI: Jaw tapers anteriorly; width about lf4-lf3

of jaw length. Fairly finely denticulated. Shank large and

pointed, flange high ly elevated.

Left MI: Jaw tapers anteriorly, fairly finely denticulated.

Basal portion skew, angular; inner wing alm ost rectangular.

Right and left MIl : Similar to the corresponding element

of the L. gracilis var. visby.

Description. - Right MI, dorsal view: Length 0 .78-1 . 1 mm,

width about 1/4-lf3 of length . The jaw tapers even ly to the

anterior (Fig. 60A:2) , ending in a fairly slender fang. The

inner margin is straight, covered by about 30 fairly small

denticles from the anteriormost part of the falcal arch,

ending slightly anterior to the deeply downfolded inner

wing.

Basal plate is not recorded.

The basal portion is of typical Lanceolatites shape, with a

large , almost triangular, sharp-ended shank, representing

about 0.2 of the jaw length. The flange is highly elevated,

smoothly rounded with a spur-like extension in the ante

rior, outer part of the basal portion. The outer margin anterior to the basal portion is smoothly undulated, with a

sickle-shaped falx.

VentraI side : Similar to L. gracilis.

Left MI, dorsal view: Length 0 .49-0.99 mm, width about

lf4-lf3 of length. The jaw tapers evenly (Fig. 60A: l ) to the anterior, ending in a fairly sJender fang. The inner margin

is straight, covered by about 30 or more fairly small denti

cles. The dentition ends before reaching the almost rectan

gular inner wing. The basal portion is irregularly rectangu

lar, with almost straight sides and a spur-like extension of

its anterior outer margin. There is a large basal furrow, and

the outer margin anterior to the basal portion is almost

crescent-shaped.

VentraI side : Similar to L. gracilis.

The right and left MIl are similar to the corresponding

element of Lanceolatites gracilis var. visby, with the following

addition: The posterior dentary of the right MIl (probably


also of the left MIl) has smaller denticles. The number of

denticles increases with increased size of the jaws. Eleven

post-cuspidal denticles have been recorded on a broken, at

least 0.88 mm long right MIl and eight denticles on a 0.40

mm long right MIl. The length of the right MIl ' s is 0.40-

0.95 mm, and of left MIl ' s 0.61-l .03 mm.

Discussion. - Lanceolatites sp. A is referred to under open

nomenclature on two gro un ds. First, the very few speci

mens encountered do not perrnit a study of the variation.

Second, the stratigraphical gap between L. gracilis and L.

sp . A makes it difficult to determine whether or not they are

of the same lineage . On the basis of the MIl morphology, a

close relation between the two seems probable.

Remarks. - A typical specimen of the taxon is the right MI

(Fig. 60A: 1 , LO 5840:6) from Strands l , Hamra Beds, unit

b, of Late Ludlow age.

Genus Langeites Kielanjaworowska 1 966

Type and only species. - Langeites glaber Kielan:Jaworowska

1 966.

Emended diagnosis. - Right MI: More or less arcuate, convex

outwards, almost . paralleI inner and outer margins,

rounded cross section. A few, wide , widely-spaced denticles

along the anterior part of the inner margin .

Left MI: Mirror image of right MI except for basal por

tion.

Right MIl: Two almost equal-sized, large , pre-cuspidal

denticles followed by a fairly large, slanting cusp. Shank

fairly short. Bight angle sharply acute , ramus long, needle

shaped.

Left MIl: Double cusp oriented to the anterior, very short

shank. Bight angle sharply acute, ramus long and needle

shaped.

Langeites gla ber Kielanjaworowska 1966 Figs. 14Q-S, 61, 62

Synonymy. - 0 1 980 Langeites glaber Kielan:Jaworowska 1 966

- Wolf, pp. 86--87, Pl. 1 0:89-94. 0 1987 Langeites glaber

Kielan:Jaworowska 1 966 - Bergman, p. 1 01-103, Figs. 1 4Q

S, 6 1 , 62.

Holotype. - No. 0.466/ 1 3b, right MI, Fig. 30:2 .

Type locality. - Unknown, erratic boulder.

Type stratum. - Ordovician or Silurian

Materialfrom Gotland. - Four right MI, 6 left MI, 2 right MIl,

2 left MIl.

Occurrence on Gotland. - Figs. 4 and 8; Middle-Late Ludlow,

Hemse Beds, Hemse Mari , SE part, and Sundre Beds. Stora

Kruse l , Vaktård 4, and Holmhilllar l .

Diagnosis. - As for the genus.

Additional description. Right MI, dorsal side : Length 0 .62-

>3. 1 mm (anteriormost part broken off and not included

in measurement) , width about 1/5 of length . The large,


Fig. 61 . Langeites glaber Kieian:Jaworowska 1 966, camera-iucida drawing of the type speeimen, right MI (Z . Pal . No. 0.466/ 1 3b) , x39.

slender jaw has almost paralleI to anteriorly slightly taper

ing sides. The cross-section is almost rounded. The inner

margin is nearly straight to pronouncedly concave with

about 1 0-16 large (wide) , anteriorly directed, knoblike

denticles, largest in the anterior part. A very short spur

exists in the middle Ludlow specimens but not in the Late

Ludlow ones.

The basal plate is rectangular and recorded in place in

the Late Ludlow specimens but not on the late Early Lud

low ones. The latter specimens have a small flange which is

almost right-angled, fairly thick, uplifted in the posterior

part and slightly downfolded along the outer margin.

Left MI, dorsal side : Length 0.89->4. 1 mm (anteriormost

and posteriormost parts broken off) , width about 1fs of

length . The large slender jaw has almost paralleI to anteri

orly slightly tapering sides. The cross-section is almost

rounded. The inner margin is almost straight to pro

nouncedly concave, with about 10 large (worn and wide) , anteriorly directed, knoblike denticles, largest on the anterior. The basal portion is almost quadratic, particularly in

the Late Ludlow specimens. A very short spur exists in the

Hemse specimens but is not seen in dorsal view on the Late

Ludlow ones. The basal furrow is pronounced, short and

fairly deep.

Right MIl, dorsal side : Length about l .5-2 .5 mm (recon

structed from two fragmented jaws) , width about half the

length . Two very large pre-cuspidal denticles of alm ost

equal size are followed by a large cusp, which is in turn

followed by fairly large denticles. The dentary with the

pre-cusps forms a weak convex arch . The shank is fairly

short with a sharply acute bight angle. The ramus is needle

shaped, long and narrow.

Left MIl, dorsal side : Length l . 55-2 .75 mm, width about

half the length . There is a large double cusp but no pro

nounced intermediate dentary. The dentary along the

inner margin is about half the jaw length . It ends conspic

uously early, leaving the posterior 0. 1 5-0.2 of the jaw un

denticulated. The bight angle is sharply aeute . The shank

is wide, short, and bluntly ended; the ramus is narrow.


Fig. 62. Langeites glaber. Specimens are in dorsal view, x1 20. From Vaktård 4, Hemse Beds, Hemse Mari SE part, 81 -35LJ; Al left MI, LO 5820: 1 ; A2 right MI, LO 5820 :2 .

Discussion. - Jaws ascribed to Langeites glaber have been

encountered in the late Early and Late Ludlow strata on

Gotland viz . Hemse Beds, Hemse Marl SE part and Sundre

Beds, middle upper part.

Due to the very limited material at hand, the relation

between the specimens from the Hemse and Sund re Beds

is somewhat dubious, though it is probable that they repre

sent the same lineage . The Late Ludlow specimens from

Gotland generally agree with Kielan-Jaworowska's description, but the late Early Ludlow jaws differ, particularly in

being almost straight to slightly convex outwards.

Remarks. - It is notable that no MIl elements have been

described from Kielan-Jaworowska's collection. The assignments here of some particular MIl elements to the same

speeies are less well-based than in reconstructions of other

Gotland apparatuses. The material derives from one local

ity (Holmhållar l ) .

The few jaws and their bad state of preservation also

explain the brevity of the description and the absenee of

variation analysis. The Holmhållar l sample was collected

by Anders Martinsson as an ostracode mari sample. It has

been processed with a coarse sieve , and the residue has

been picked over for ostracodes. I have only had access to

the picked out scolecodonts, and so do not know if these

are representative for the sample .

Silurian paulinitid polychaetes from Gotland 1 1 3

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Wolf, G. 1976: Bau undFunktion der Kiejerorgane von Polychaeten. [ On the morphology and function of polychaete jaws. ] 55 pp. Zoologisches Institut und Museum, Universitat Hamburg.

Wolf, G. 1 980: Morphologische Untersuchungen an den Kieferapparaten einiger rezenter und fossiler Eunicoidea (Polychaeta) . Senckenbergiana maritima 12:1/4, 1-182 .

Zebera, K. 1 935: Les Conodontes et les Scolecodontes du Barrandien. Bulletin international de l'Academie des Sciences de Boheme 36, 88-96.

FOSSILS AND STRATA 25 ( 1 989) Silunan paulinitid polychaetes from Gotland 1 1 9

Appendix: Localities In the list of references to each locality, an asterisk marks the publication in which a major description of the locality is found. Other references are given only if not included in 'Reference localities for paJaeontology and geology in the Silurian of Gotland' (Laufeld 1 974b) . The grid references refer to the Swedish National Grid system measured from the 2nd topographical map edition and (within parentheses) the UTM system from the 1 st edition.

Silicified Fossils from Gotland' marked PSSFG) and some further samples from the Swedish Museum of Natural History, Stockholm,

Ken t Larsson, Lund, Anders Martinsson, Uppsala, Doris Fredholm, Lund, Sven Laufeld, Uppsala, and Sven Stridsberg, Lund.

The neutral term 'Beds' has been used for designating Hede 's major stratigraphical units.

Only part of the jawed annelid fauna ( the paulinitids) is described in this paper. Paulinitids are reported by specific name, all other species of jawed annelid as ' annelid jaws ' . If only fragments and no identifiable annelid remains are found, 'fragments' is noted.

The sample codes con sist of the two last figures of the sampling year, a sample number for thatyear, and the initials ofthe collector. Apart from my own samples I have received a large num ber from LennartJeppsson (including a few samples from 'Project Silurian

AJ M U N D E 1 . Hemse beds , Hemse Mari NW part. Age: LudJow, probably Bringewoodian. References:Laufeld 1 974a, b*; Laufeld & Jeppsson 1 976; Larsson 1 979; Jeppsson 1 983; Ramsk61d 1 985; Frykman 1 989, Samples/jawedannelids: 7 1 -40LJ, bottom of the ditch, LanceolaUles gracilis, Kettnerites (K.) cf. martinssonii, K. (A.) sisyphi, K. (K.) cf. burgensis, annelid jaws; 75-77CB, bottom of the ditch , fragments; 77-36LJ, bottom of the ditch, few annelid jaws.

AJSTU DDEN 1 . 639599 1 67863 (CJ 6866 9246), ca 4250 m SSE of Boge church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 69 Slite.

Shore exposure comprising about 1 00-1 50 m north of the small point along the shore and some tens of meters out to sea. Slite Beds, Slite Mari, probably slightly older than Pentamerus gothlandicus. Age: Wenlock. Samplesljawed annelids: 83-6CB, fossiliferous grey calcarenite immediately below the mean water level, Kettnerites (A) sisyphi, Gotlandites sfitensis, annelid jaws; 83-7GB, grey dense calcarenite about 1 0 m beyond the shorel ine and 0.2 m below the mean water level covered by a few m m of quaternary sediment, Gotlandites slitensis, other paul i nitid jaws, annelid jaws.

ALBY 1 . Slite Beds, Slite MarI. Age: Wenlock. References: Laufeld 1 974b*; Larsson 1 979. Samplesljawedannelids: 67-20LJ, Kettnerites (A) sisyphi, annelid jaws.

ALBY 2. Slite Beds, Slite MarI. Age: Wenlock. References: Laufeld 1 974a, b*; Ramskold 1 983. Samplesljawed annelids: 75-1 7CB, excavated material, annelid jaws.

ALBY 4. Slite Beds, Slite MarI. Age: Wenlock. References: Laufeld 1 974a, b*. Samples/jawed annelids: 77- 1 6GB, uppermost bed about 0.75 m below ground level, anneJid jaws.

AMLINGS 1 . Hemse Beds, Hemse Mari NW part. Age: Ludlow; probably Late Bringewoodian, possibly earliest Leintwardinian. References:Laufeld 1 974a, b*; Laufeld & Jeppsson 1 976; Larsson 1 979; RamskOld 1 983; Jeppsson 1 983*; Fredholm 1 988. Sampfesljawed annelids: 7 1 - 1 1 1 LJ, top of the section, Kettnerites (K.) huberti, K (A) microdentatus, annelid jaws; 75-73CB, 0. 1 0 m above the bottom of the ditch, same sample level as 66-1 70SL (Laufeld 1 974a), Kettnerites (Kl pofonensis, K. (A) sisyphi aff. kJasaardensis, K. (A) microdentatus, annelid jaws; 75-74CB, 0 . 1 5 m above the bottom of the ditch , same level as sample G66-1 70SL (Laufeld 1 974a), Lanceolatites graeilis, Kettnerites(K) huberti, K. (A) sisyphi, annelid jaws; 77-39PSSFG, Kettnerites (K) martinssonii, K. (K.) cf. burgensis, annelid jaws.

ANGMANS 2. Hemse Beds, unit b. Age: Ludlow, Eltonian. Reference: Fredholm 1 988*. Samples/jawedannelids: 83-50LJ, excavated material, Kettnerites (K.) bankvaetensis, K (K) huberti, annelid jaws.

ANSARVE 1 . Hogklint Beds, southwestern facies, upper part. Age: Early WenJock. References: Laufeld 1 974b*, Larsson 1 979. Samples/jawedannelids: 79-46LJ, bottom of the ditch , Ketlnerites (K.) mar1inssonii, K. (A) sisyphi, K (K.l abraham isaac, K (Al sisyphivar. valle, Hindenites angustus, annelid jaws.

AR 1 . HogkJint Beds, upper part. Age: Early Wenlock. Reference: Larsson 1 979*. SampfesljawedanneJids:84-13U 0.25 to 0.5 m above base of the section, Kettnerites (K) martinssonii, K. (K.) abraham isaac, Hindenites cf. gladiatus, annelid jaws.

ASK RYGGEN 2 . Slite Beds, Slite Siltstone, top. Age: Wenlock. References: Laufeld 1 974a, b*; Bergman 1 980a. Samplesljawedannefids: 7 1 -S2LJ, no annelid jaws.

AURSVIKEN 1 . HOgklint Beds, lower-middle part. Age:Early Wenlock. References: Laufeld 1 974a, b*. Sampfesljawed annelids: 77-25GB, surface exposure, Kettnerites (K) martinssonii, K. (A) microdentatus, Hindenites cf. gladiatus, annelid jaws.

AUTSARVE 1 . Hemse Mari NW part. Age: Early Middle Ludlow. References: Laufeld 1 974a, b*, Jeppsson 1 983'; Fredholm 1 988; Frykman 1 989.

Sampfeljawed annelids: 7 1 -43LJ , surface exposure, Lanceolatites graciJis, Kettnerites {K.) martinssonii, K (K.) huberti, K. (A) sisyph,; annelid jaws; 75-60GB, loose slab, no annelid jaws.

BAJU 1 . 637823 1 67773 (CJ 6639 7480), ca 4 1 00 m ENE of Anga church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 70 Katthammarsvik.

Low cliff at the shore at the southernmost tip of the point about 50 m south of the pier. Kfinteberg Beds{?). Age: Late Wenlock - Early Ludlow. Samplesljawedannelids: 78-1 2CB, 0. 1 5 m below top of the exposure, Kettnerites {K) cf. bankvaetensis, K. (K.) mar1inssonii, K (K) polonensis, annelid jaws.

BAN DLUNDE 1 . Eke Beds, lower part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO. SampJesljawed annelids: 76- 1 2GB, loose slab from a temporary ditch excavation, Kettnerites (K) martinssonii, K (K) cf. huberti, annelid jaws.

BANKVAT 1 . Hamra Beds, unit b. Age: Latest Ludlow. References: Lauleld 1 974a, bO; Larsson 1 979; Jeppsson 1 982, 1 983. Samplesljawedannefids:81 -39LJ, some 1 0 m N of the lake (existing only during wet periods) and slightJy south of sample location of 82-33CB, Kettnerites (K) bankvaetensis, K (K) huberti, annelid jaws; Samples 82-27GB to 32G B : Aboul 1 55 m N E of the stone fence and 1 0-1 5 m N ol the beach, i n a shallow shore exposure, 82-27CB, bed 0 . 1 2-0.20 m below the uppermost Sil urian rock surface, undulating, competent, brownish-grey calcarenite between soft calcilutite, Kettnerites (K) bankvaetensis, annelid jaws; 82-28CB, 20-50 mm thick bed and about 0 . 1 0 m below sulface, Kettnerites (K) bankvaetensis, K. (K) huberti, annelid jaws; 82-29GB, 1 0-30 mm thick bed and about 0.08 m below surface, Kettnerites (K) bankvaetensis, annelid jaws; 82-30GB, 30-40 mm thick bed about 0.05 m below surface, Kettnerites (K.) bankvaetensis. K (K) huberti, annelid jaws; 82-3 1 CB, 1 0-30 mm thick bed and aoout 0 .03 m below surface, Kettnerites (K.) bankvaetensis, annelid jaws; 82-32GB, about 30 mm thick, comprising the uppermost bed, Kettnerites (K) bankvaetensis, K. (K.) huberti, annelid jaws; 82-33GB, about 1 5 0 m NW of samples 82-27CB to 82-32GB and 1 00 m SE of the 82-34GB, about 0.30-0.40 m below the soi l surface, Kettnerites (K.) bankvaetensis, K (K.) huberti, annelid jaws; 82-34CB, about 250 m NW of the beach parallei to and 1 00 m N E of the stone fence, 0 . 1 m below ground surface, Kettnerites (K) bankvaetensis, K. (K.) huber1i, annelid jaws; 83- 1 5GB, ca 50 m E of the small ruin adjacent to the stone fence, the second erratic boulder from the ruin, about 1 m in diameter, sample from the stromatoporoid-rich bed, the topmost bed, Kettnerites (K) bankvaetensis, annelid jaws; 83- 1 6GB, calcarenite with calcareous algae, 0 . 1 5-0.20 m below surface, Kettnerites (K.) bankvaetensis, annelid jaws; 83-1 7GB, ca 300 m N of the ruin and 75 m E of the stone fence, 0.25 m below soil surface, ca\carenitic nodules in soft calcilutite, Lanceofatites sp. A, Kettnerites (K) bankvaetensis. K. (K) pofonensis, annelid jaws.

BARA 1 . Slite Beds, Halla Beds, oolite. Age: Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 982, 1 983; Ramsk61d 1 983; Frykman 1 989. Samplesljawed annelids: Slite Bads: 70-1 2LJ, on the slape above the quarry, no annelid jaws; Halla Beds: 67-24LJ, in the quarry, no annelid jaws; 78-4CB, 0.75 m above the bottom i n the section, no annelid jaws; 70- 1 3LJ, on the slope near the telephone pole above the quarry, no annelid jaws.

BARKARVEARD 1 . Hamra Beds, unit c. Age: Late Ludlow. References: Laufeld 1 974a, b*; Larsson 1 979. Sampfesljawed annelids: 76- 1 4CB, shore exposure, Kettnerites (K) huber1i, K. (K.) polonensis, annelid jaws.

BARSHAG EUDD 2. Hamra Beds, unit c. Age: Late Ludlow. References: Laufeld 1 974a, bO. Sampfes/jawed annelids: 76-26GB, shore exposure, pauli nitid frag-ment, annelid jaws.

BATA 1 . Slite Beds, unit f (Rhipidium tenuistriatum Beds) . Age: Early Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979; Jeppsson 1 983. Samplesljawed annelids: 77-27CB, 0. 1 5 m below the uppermost bed in the ditch, no annelid jaws.

BATELS 1 . Klinteberg Beds, unit c . Age: Late Wenlock. References: Laufeld 1 974a, b*; Jeppsson 1 983; Frykman 1 989.

Samplesljawed annelids: 7 1 -89LJ, the lowest exposed bed, no annelid jaws; 71 -90LJ, 0 .90 m above the lowest exposed bed, n o annelid jaws; 7 1 -91 LJ , 2 . 1 0 m above the base o f t h e section, t o p o f t h e section, no annelid jaws.

BATELS 2 . Klinteberg Beds, unit b. Age: Late Wenlock. References: Laufeld 1 974a, bO; Frykman 1 989. Samplesljawedannelids: 7 1 -92LJ , 10p of the section, no annelid jaws.

BINGERS KVARN 2. Tofta Beds. Age: Early Wenlock. References: Laufeld 1 974a, b·. Samplesljawed annelids: 75-27CB, uppermost bed of the road exposure, no annelid jaws.

BJARGES 1 . Mulde Beds, upper part. Age: Late Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979; Frykman 1 989. Samplesljawed anneJids: 7 1 -9LJ, excavated material, Kettnerites (A) microdentatus, paul in itid sp. , annelid jaws; 75-68CB, about 0.5 m below ground level, Kettnerites (A) sisyphi, annelid jaws.

BJARGES 2. 635275 1 64223 (CJ 29 1 1 521 4), ca 2725 m WSW of Eksta church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 64 Hemse.

Ditch exposure north of and along the private road, where a pipe u nderlies the road, about 350 m SW of Ajvide and about 1 75 m W of the small houses lo the left of the H in Hagur. Mulde Beds, upper part. Age: Late Wenlock. Samples/anneJid jaws: 79- 1 1 LJ, bottom of the ditch, Kettnerites (A) sisyphi, annelid jaws.

BLAHALL 1 . Mulde Beds, lower part. Age: Late Wenlock. References: Laufald 1 974a, bO; Glaesson 1 979; Larsson 1 979; Poulsen et al. 1 982; Jeppsson 1 983; Ramskold 1 984, 1 985; Fryk-man 1 989. Reference level: About 2 m above water level, the re is a protruding bed that can be lollowed for at least 50-1 00 m . The lower surface of this bed is designated as reference leveL Samplesljawedannelids: 7 1 - 1 40LJ, 1 . 50 m below reference level, Kettnerites (K) martinssonii var. mulde, K. (A) sisyphi, annelid jaws; 75-65CB, 0.50 m below the top of the section, Kettnerites {K) martinssonii var. mulde, K (A) sisyphi, annelid jaws; 75-66GB, 1 .2 m below the top of the section, Kettnerites (K) martinssoniivar. mulde, K. (A) sisyphi, annelid jaws.

BODUDD 1 . Hemse Beds, Hemse Mari uppermost part and Eke Beds, basal part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO, c ; Larsson 1 979; Jeppsson 1 982, 1 983'; Cherns 1 983"; Fredholm 1 988. Samplesljawed annefids: Hamse Beds, uppermost part: 75-t OOCS, just above mean water level , Kettnerites (A) sisyphi kJasaardensis, annelid jaws; 8 1 -30LJ, in the southeaslernmost part of the exposure, at mean water level, in the water, the lowest bed about 5 m NW of the sea-wall, annelid jaws, 81 -3 1 LJ, about 0.3 m higher than 8 1 -30, 2 m NW of the sea-watl, fragments; 8 1 -32LJ, topmost Hemse Beds, Kettnerites (A) sisyphiklasaardensis, annelid jaws; Iowast Eke Beds: 8 1 -33LJ, about 1 00 m NW of the tail of erratic boulders in the sea and 0.3 m below the uppermost exposed bed, Kettnerites (Kl martinssonii, K. (A) sisyphi klasaardensis, K (Kl huberti, K (A) cf. microdentatus, annelid jaws; 8 1 -34LJ , the uppermost exposed bed, Lanceolatites gracilis var. visby, Kettnerites (A) sisyphi klasaardensis, annelid jaws.

BODUDD 2. Eke Beds, lowest part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979. Samples/jawed annelids: 75-1 0 1 GB, from the shoreline, Lanceolatites gracilis, Kettnerites (K) huberti, annelid jaws.

BODUDD 3 . Hemse Beds, Hemse Mari uppermost part. Age: Ludlow, Late Leintwardinian. References: Hede 1 9 1 9:21 , Loc. 9 ; Hede 1 942:20, Loe 1 C ; Jeppsson 1 983'. Samplesljawedannelids: 7 1 - 1 5 1 LJ, shallow shore exposure available at low water, Lanceolatites gracilis, Kettnerites (K.) martinssonN, K (A) sisyphi klasaardensis, K. (K.) huberti, K (K.l cf. burgensis, annelid jaws.

BOFR I D E 1 . Klinteberg Beds, upper part. Age: Late Wenlock to , Early Ludlow.

References: Laufeld 1 974a, b*; Larsson 1 979; Frykman 1 989. Samplesljawed annelids: 70-32LJ, 0.0-0 . 1 m above the lower marly bed, Kettnerites (K) cf. bankvaetensis, annelid jaws; 75-70GB, 1 . 25 m .below reference level, Kettnerites (K) cf. martins-


sonii, K. (A.) sisyphi cf. var. valle, annetid jaws; 75-7 1 CS, 2.0 m below reference level, Kettnerites (K.l cf. bankvaetensis, annelid jaws; 75·72CB, 1 .0 m above reference level. no annelid jaws.

BOFRIDE 2. Klinteberg Beds, upper part. Age: Early Ludlow. Referenees: Laufeld 1 974a, b*; Frykman 1 989. Samples/jawed annelids: 75-69CB, 1 . 5 m from top of the section, no annelid jaws.

BOTTAAVE 1 . Hamra Beds, unit b. Age: Ludlow, Whitcliffian. References: Laufeld 1 974a, b'; Laufeld & Jeppsson 1 976. Samples/jawed annelids: 77-32LJ, shallow ditch, KeNnerites (K.l bankvaetensis, annelid jaws; 83-21 CB, shallow ditch, paul in itid fragments, annelid jaws.

BOTTARVE 2. Hamra Bads, unit b. Age: Ludlow, Whitcliffian. References: Laufeld 1 974a, b'; Laufeld & Jeppsson 1 976. Samp/esljawed annefids: 76-21 GB, excavated material, Kettneriles (Kl cf. bankvaetensis, paulinitid fragments, annelid jaws; 76-22GB, excavated material, Kettnerites (K.l cf. bankvaetensis, annelid jaws; 63- 1 9GB, ca . 20 m W of the sharp bend of the ditch which continues i n an E-W direction, botlom bed, Kettnerites (K.l martinssonii, K. (A) microdentatus, annalid jaws; 83-20GB, immediataly above 83- 1 9GB, pau l initid fragments.

BOTVALDE , . Klinteberg Beds, unit e. Age:Wenlock, Late Homerian. Reference:Jeppsson 1 983'; Frykman 1 989. Samp/esljawedannefids: 75-37LJ, small quarry, annelid jaws.

BOTVIDE 1 . Hemse Mari uppermost part and Eke Beds basal part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, b'; Jeppsson 1 974, p. 1 0 , 1 982, 1 983; Larsson 1 979; Laufeld & Martinsson 1 98 1 ; Gharns 1 982*, 1 983·. Sampfesljawed annelids: Hemse Beds: 69- 1 8LJ, 0.60-0.70 m below the conglomerate level, Kettnerites (K.) cf. pofonensis, K. (K.) cf. burgensis; 69- 1 9LJ, 0 .05-0 . 1 0 m below the conglomerate level, paulinitid sp. 1M I, annelid jaws; 69-20LJ, 0.0-0.05 below the conglomerate level, Kettnerites (K.l huberti, annelid jaws; Eke Beds: 69-22LJ, 0.0-0.05 above the conglomerate level, Kettnerites (Kl huberti, K (Kl burgensis; 69-23LJ, 0 . 1 0-0. 1 5 m above the conglomerate level, no annelid jaws; 69-24LJ, 0.25 above the conglomerate level, no annelid jaws; 69-25LJ, on the northern slope 0.5 m below tap of the isolated sea stack, no annelid jaws.

BRINGES 1 . Sundre Beds, lower part. Age: Latest Ludlow. References: Laufeld 1 974a, b·. Samp/esljawedannelids: 7 1 - 1 86LJ, no annelid jaws.

BRINGES 3 . 632 1 95 1 65562 ICJ 4006 2043), ca 4020 m N E of Hamra church. Topographical map sheet 5 I Hoburgen SD & 5 J Hemse SV. Geological map sheet Aa 1 52 Burgsvik.

Abandoned, shallow quarry, 400 m SSE of the manor house at Bringes. Sundre Beds, lower part. Age: Latest Ludlow. Sampfesljawedannefids :7 1 - 1 87a LJ, Kettnerites (K) cf. bankvaetensis, annelid jaws.

BROA 2. Slite Beds, Ryssnas limestone. Age: Wenlock, middle part. Referenees: Laufeld 1 974a, b* Samp/esljawed annefids: 78- 1 6GB, sample from the reference level, Kettneriles (K) jacobi, K (A) sisyphi, annelid jaws.

BROTRASKKROKEN ,. Hemse Beds, midd le upper part. Age: Ludlow. References: Laufeld 1 974a, b'; Larsson 1 979. Samp/esljawedannefids:71 -44LJ , lowest exposed bed, no annelid jaws.

BURGEN , . Burgsvik Beds, upper part. Age:late Ludlow, possibly Whitcliffian. References: laufeld 1 974a, bO; Larsson 1 979. Samplesljawed annefid jaws: 82-21 GB, about 0.85 m below the boundary to the crinoid-rich beds, conglomeratic balls, no annelid jaws; 82-22GB, from the same conglomeratic level as the former sample but also including the oolitic limestone which embeds the bal ls , ca 0.85 m below the bed rich in crinoids, pauli nitid fragments; 82-23GB, immediately below the crinoid-rich bed about 2.35 above the botlom of the quarry, no annelid jaws; 82-24GB, probably Hamra Beds, the lowest crinoid-rich layer about 2.40 m above the quarry floor, no annelid jaws.

BURGEN 5. Eke Beds, lower part. Age: ludlow, possibly Late Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979. Samplesljawed annelids: 77-32CB, ditch seclion, brownish bluegrey calcibiorudite, no annelid jaws.

BURGEN 9. Burgsvik Beds, Burgsvik Oolite. Age: Late Ludlow. Referenees: Larsson 1 979' Sampfesljawed annelids: 7 1 - 1 32LJ , northern wal l , low in the section, no annelid jaws; 7 1 - 1 33LJ, 3-4 m above 7 1 - 1 32, upper part of the seclion, no annelid jaws.

BUSKE 1. Lower Visby Beds, Upper Visby Beds. Age: Early Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979; Odin et al. 1 984*. Auxiliary reference leve/: The boundary between the Lower and Upper Visby Beds has been reported at 0 . 75 m above the base of the section (Hede 1 940 : 1 3, Martinsson 1 962:47, Laufeld 1 974b), but according to Lennart Jeppsson (personal communicalion, 1 986) and Odin et al. (1 984) it is aboul 2.8 m above the base. Martinsson's level cannot be identified with the necessary accuracy, thus it has not been used as reference level by Lennart Jeppsson. Instead, the sampled levels have been measured from the best visible bentonite, which is about 1 .6 m above the base of

the section and about 1 .2 m below the level abundant in large solitary rugose corals. Samples/jawedannefids: Lower Visby Beds, unit e: 79-4OW , 1 .3-1 . 1 m below the auxiliary reference level, Kettnerites (K.) versabilis form A, B, and C, Laneeo/atites gracilis, Kettnerites (K.) martinssonii, K (A) cf. mierodentatus, K (Kl abraham abraham, annelid jaws; 79-41 LJ, 0.2-0.25 m above the auxil iary reference level, Kettnerites (K) versabiJis form A and B, K (K) martinssonii, K (K) abraham abraham, annelid jaws.

DACKER , . Slite Beds, unit e. Age:Wenlock. References: Laufeld 1 974b' ; Glaesson 1 979. Sampfesljawedannefids: 67- 1 4LJ, top of the section, no annelid jaws.

DAPPS l . Mulde Beds, upper part. Age: Late Wenlock. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Frykman 1 989. Samp/es/jawedannelids: 81 ·56U, about 15 m SE of the road, l m section exposed, sample from the 0 . 1 5 m thick bed, Kettnerites (Kl bankvaetensis, other paulinitid taxa, annelid jaws; 82-45GB, 1 .48-1 .55 m below the surface of the SW corner of a concrete ramp, built for the former railway, Kettnerites (K.) martinssonii var. mulde K. (A) sisyphi sisyphi, annelid jaws.

DAPPS 2. Mulde Beds, upper part. Age: Late Wenlock. Referenees: Laufeld 1 974a, b'; Larsson 1 979. Samples/jawed annefids: 75-25GB, 0.5 m below the tap of the section, Kettnerites (K) martinssonii, K (A) sisyphi sisyphi, annelid jaws; 82-37G8, top of the section, 20-30 m m thick bed, Hindenites g/adiatus, Kettnerites (Kl martinssonii, K. (A) sisyphi sisyphi, annelid jaws; 82-38GB, 0 .60 m below the top of the section and 1 .0 m above railway embankment, about 50 mm thick bed composed of bluish-grey, dense l imestone, Kettnerites (K.) martinssonii, K. (A) sisyphi sisyphi, annelid jaws; 82-39CB, 30 m north and about 1 .70 above the railway embankment, Kettnerites (Kl martinssonii, K. (A) sisyphisisyphi, annelid jaws, 82-40CB, 0.98 m below the lower surface of 82-39GB, Kettnerites (Kl martinssonii, K. (Al sisyphi sisyphi, annelid jaws.

DIGRANS 1 . Hamra Beds, lower part. Age: Ludlow, Whitcliffian. References: Laufeld 1 974a, b' Samples/jawed annefids: 75- 1 5LJ, ditch seclion, pauli nitid jaw, annelid jaws.

DJAU PVIKSUDDEN 4. 637500 1 67953 ICJ 6796 7 1 49), ca 6040 m NW of Ostergarn church. Topographical map sheet 6 J Roma SD. Geological map sheet Aa 1 70 Katlhammarsvik.

About 1 00 m N E of Djaupviksudden 1 (described i n the geologicat map description). Hemse Bads, unit b and c. Age: Ludlow. Samplesljawedannelids: 7 1 - 1 55LJ, 0-1 0 mm, immediately above the corroded surface, annelid jaws; 7 1 - 1 57LJ, from the lower part of the three levels in the section, described in Hede's 1 960 guide, paul in itid fragments, annelid jaws; 7 1 - 1 59LJ, from the uppermost level , Kettnerites (K) huberti, K. (K) p% nensis, annelid jaws.

DJUPVIK 1 . Mulde Beds, lower part. Age: Late Wenlock. Referenees: Laufeld 1 974a, bO; Larsson 1 979, Jeppsson 1 982, 1 983; Odin et al. 1 984, 1 986. Samples/jawed annelids: 75-67GB, middle part ol the section, Kettnerites (K) martinssonii var. mulde, K (A) sisyphi sisyphi, annelid jaws; 84-21 LJ, the calcarenitic bed between the bentonite bed selected as reference level by Jeppsson ( 1 983:1 40) and the bentonite 0.2 m above it, Kettnerites (K) martinssonii, K. (A) sisyphi sisyphi, annelid jaws.

DJUPVIK 2. Mulde Beds, lower part. Age: Late Wenlock. Reference:Jeppsson 1 983*; Odin at al . 1 984'. Reference /evel: 1 .05 m above the distinct mari horizon (erroneously given as 1 .50 m in Jeppsson 1 963 ; l . Jeppsson, personal communication, 1 986). Samp/esljawedannelids: 70-25LJ, 0 .0-0.2 m above the reference level, Kettnerites (A) sisyphi sisyphi, annelid jaws; 70-26LJ, 1 .05-0.95 m below the reference level, Kettnerites (K) martinssonii, K. (A.) sisyphi sisyphi, annelid jaws; 70-27LJ, 1 .07-1 .20 m above the reference level, Kettnerites cf. (K.) martinssonii, K (A l sisyphi sisyphi, anne lid jaws; 70-28LJ, 2.24 m above the reference level, top of the section, Kettnerites (K.) cf. martinssonii, K (A) sisyphi sisyphi, annelid jaws; 70-29LJ, 1 .95-2.05 m below the reference level, Kettnerites (K) martinssonii, K. (A) cf. sisyphi, annelid jaws.

DJUPVIK 3. 635577 1 64096 ICJ 2804 5525), ca 4275 m SW of Frejel church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 64 Hemse.

From the solitary house west of the road to the fau k, about 50 m SW ol the house. Djupvik 1 starts beyond the fault. A section of about 7.2 m Med strata is exposed. Mulde Beds. Age: Late Wenlock. Samples/jawed annelids: 84- 1 8LJ, the lowest exposed bed at low water, Kettnerites (K) martinssonii mulde type, K (A) sisyphi sisyphi, annelid jaws; 84- 1 9LJ, the uppermost exposed bed, Kettnerites (K) martinssonii mulde type, annelid jaws.

DJUPVIK 4. 635576 1 64 1 06 ICJ 281 3 5523), ca 4425 WNW of Eksta church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 64 Hemse.

Rivulet section in the gully just N E of the solitary house west of the road and up stream some ten metres beyond the road. Mulde Beds. Age: Late Wenlock. Samples/jawed annelids: 84-20LJ, uppermost bed in the section, Kettnerites (K.) martinssonii mulde type, annelid jaws.

DRAKARVE , . Hemse Bads, Hemse Mari, uppermost part. Age: Ludlow, probably Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979.


Samples/jawed annelids: 75-99GB, about 0 . 5 m below t h e soi l surface and 0.7 m above the base of the normally water lilIed holtow, no annelid jaws.

ENHOLMEN 2. 640083 1 6800 1 ICJ 7037 9722), ca 21 00 m S E of Slite church. Topographical map sheet 7 J Fåresund SO & NO. Geological map sheet Aa 1 69 Slite.

Shore exposure on the N E side of the Enholmen island. Slite Beds, unit g . Age: Wenlock. Referenees: Hede 1 928, p. 4 1 , l ines 24-26; Martinsson 1 962, p . 52 (references to Enholmen island i n general). Samp/es/jawed annelids: 75-9GB, shore exposure, no annelid jaws.

FÅGELHAMMAR 3. 6361 73 1 67652 ICJ 6392 5847), ca 7550 m SW of Gammelgarn church. Topographical map sheet 6 J Roma SO. Geological map sheet Aa 1 70 Katthammarsvik.

Area with loose slabs, SW of the sea stad<. field, SSW of Fågelhammar 1 , about 50 m SW of the triangulation point at Folhammar. Hemse Beds, upper part. Age: Ludlow. Referenees: Hede 1 929, p . 53, l ine 1 7 , to p . 54, l ine 4. Samp/es/jawedannelids:71 -21 8LJ, loose conglomeratic flat slabs, some 10 m SW of the southernmost sea stack on the beach, Laneeo/atites gracilis, Kettnerites (K.) polonensis, annelid jaws.

FAKLE , . Hemse Beds, unit c. Age: Ludlow, probably Bringewoodian. References: Laufeld 1 974a, bO; Larsson 1 979; Fredholm 1 988. Samplesljawed annelids: 7 1 - 1 62LJ, the lowest brownish bed, Ketfnerites (Kl pofonensis, pau l in itid fragments, annelid jaws.

FALUDDEN , . Sundre Beds, lower part. Age: Latest Ludlow. Referenees: Laufeld 1 974a, b'; Larsson 1 979. Samp/es/jawed annelids: 76- 1 5GB, uppermost bed between 5tromatoporoids, no annelid jaws.

FALUDDEN 2. Hamra Beds and Su ndre Beds. Age: Latest Ludlow. References: Laufeld 1 974a, b' Samp/esljawedannelids: Hamra Beds, unit c: 76-1 6CB, uppermost bed, Kettnerites (K) bankvaetensis, K (K) huberti, K (Kl p% nensis, annelid jaws.

FARDUME 1. 641 275 1 68555 ICK 7681 086 1 ) , ca 3650 m S of Rute church. Topographical map sheet 7 J Fåresund SD & NO. Geotogical map sheet Aa 171 Kappelshamn.

Quarry along the road about 1 00-1 50 m NNW of Fardume ruin. Slite Beds, unit g . Age:Wenlock. Reference: Hede 1 933:44. Samplesljawed annefids: 84-1 7LJ, 1 .5 m above quarry floar, thin l imestone slabs in a deeply eroded lutitic bed, Gotlanditesslitensis, Kettnerites (K.) martinssonii, K. (K.) p% nensis var. gandarve, K (A) cf. sisyphi, Hindenites gladiatus, annelid jaws.

FÅRO SKOLA 1 . Slite Beds, Slite MarI. Age: Wenlock. Referenees: Laufeld 1 974b'; Larsson 1 979. Samp/esljawed annelids: 78- 1 5CB, uppermost bed, Kettnerites (Al sisyphi sisyphi, annelid jaws.

FIE 3. Hemse Beds, Hemse Mari S E part. Age: Ludlow, Leintwardinian(?). Reference: Jeppsson 1 983; Fredholm 1 988. Samplesljawedannelids:71 - 1 28LJ, botlom of the ditch, Kettnerites (K.) bankvaetensis, K. (K) martinssonii, K (K) huberti, K (K.) cf. burgensis, annelid jaws.

FJALE 2. Klinteberg Beds, unit d . Age: Latest Wenlock - earliest Ludlow. References: Laufeld 1 974a, bO; Frykman 1 989. Samp/esljawed annelids: 77-6GB, shallow ditch , 0.25 m below ground level, Kettnerites (K) bankvaetensis, annelid jaws

FJ.AlE 3. Klinteberg Beds, unit e . Age: Latest Wenlock - earliest Ludlow. References: Laufeld 1 974a, b'; Frykman 1 989. Samp/es/jawed annelids: 73-31 LJ , from the 0.3 m exposed, fragments; 77-7GB, 0 .50 m below ground level , Lanceolatites gracilis, L . gracilis var. visby, Kettnerites (K) bankvaetensis, K (K.) martinssonii, K (A.l fjaelensis, Hindenites sp. , annelid jaws.

FJARDINGE 1 . Klinteberg Beds, unit b. Age: Latest Wenlock -earliest Ludlow. References: Laufeld 1 974a, b'; Larsson 1 979; Frykman 1 989. Samplesljawed annefids: 75-33LJ, top of accessible seclion, no annelid jaws; 75-34LJ, lowest accessible bed aboul 1 m below 75-33, Kettnerites (K) cf. bankvaetensis, K (K.) cf. pofonensis, annelid jaws; 77-4GB, uppermost bed of the section, no annelid jaws; 77-5GB, loose slabs probably excavated from the ditch, Kettnerites (K.) bankvaetensis, K (Kl martinssonii, annelid jaws.

FOLLINGBO 2. Slite Beds, Slite Mari, NW part. Age: Wenlock. References: Laufeld 1 974a, bO ; Larsson 1 979. Samplesljawed annelids: 75-1 OGB, sample from the botlom of the ditch, Kettnerites (K.) martinssonii, K (A) mierodentatus, annetid jaws.

FOLLING BO 3. Slite Beds, Slite Mari, NW part. Age: Wenlock. References: Laufeld 1 974a, b·; Larsson 1 979. Sampfes/jawedannelids: 71 -68LJ, lowest accessible bed, Kettnerites (A) sisyphi, fragments.

FOLLINGBO 4. Slite Beds, unit g . Age: Wenlock. Referenees: Laufeld 1 974a, bO; Glaesson 1 979; Larsson 1 979. Samp/esljawed annelids: 71 -71 LJ, 0.5 m below the road surface, fragments.


FOlLiNGBO 1 2. 638848 1 65401 I C J 4353 8683), ca 1 430 N of Follingbo church. Topographical map sheet 6 J Visby NV & NO. Geological Map sheet Aa 1 83 Visby & Lurnrnelunda.

Temporary excavation about 50 m west of the almast N-8 directed road (the road between the main roads Visby-Endre and Visby-Roma) and some 1 0 m lett of the smaU road to the sanatorium, at the first right turn on the small road. Slite Bads, Slite Mari, northwesternmost part. Age: Wenlock. Samplesljawed annelids: 80-1 GB, loose slab from excavation, Kettnerites (K.) martinssonii, Hindenites angustus, annelid jaws.

FROJEL 2. Mulde Beds, uppermost part. Age: Late Wenlock. References: Laufeld 1 974a, b*; Frykman 1 989. Samples/jawed annelids: 70-30LJ, shallow ditch, paulinitid fragments, annelid jaws.

GALGBERGET 1 . Hogklint, unit c, and Tofta Beds. Age: Early Wenlock. References: Laufeld 1974a, b*. Samplesljawedannefids: 67-4LJ, lowermost Tofta bed, fragments; 67-5LJ , uppermost Hogklint, unit c, no annelid jaws.

GAMLA HAMN 1 . Hogklint Beds, unit b. Age: Early Wenlock. Reference: Laufeld 1974b*. Samplesljawedannelids: 7?-23CB O.5 m below the uppermost part of the stratified bed between the sea stacks, Kettnerites (K.) martlnssonii, anne l id jaws.

GAMLA HAMN 2. Hogklint Beds, unit c. Age: Early Wenlock. Reference: Laufeld 1 974b*. Samplesljawed annelids: Hogklint Beds, unit c : 67- 1 8LJ, annelid jaws; Slite 8eds, unit c: 67- 1 9LJ, about 2 m above 67·1 8, no annel id jaws.

GANDARVE 1 . Halla Beds. Age: Late Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979. Samplesljawed annelids: 7 1 -81 LJ, Hindenites gladiatus, Kettnerites (K.) martinssonii, K (K) polonensis var. gandarve, K (A) sisyphi sisyphi, K (A) sisyphi var. val le, annelid jaws; 75- 1 3CB, base of the section, no annelid jaws.

GANDARVE 2. Halla Beds. Age: Late Wenlock. References: Laufeld 1 974a, b*. Sampfesljawed anne/ids: 75- 1 2CB, uppermost prominent calciruditie bed, fragments; 84-1 1 3CB, third bed from the top in the section, Kettnerites (K) martinssonii, K (A) sisyphi sisyphi, annel id jaws.

GANE 2. Slite Beds, Slite MarI. Age: Wenlock. References: Hede 1 92B·, 1 942 10cality 2 1 ; Larsson 1 979*; Ramskold 1 983. Samplesljawed annelids: 83-29 U , 0.5 m exposed, sample from the middle, eroded part, 200 m SW of the private road along the former railway l ine, Kettnerites (K) bankvaetensis, annelid jaws.

GAN NES 2. Hemse Beds, Hemse Mari, unit c o r d (?). Age:Ludlow, Brlngewoodian or Early Leintwardinian. References: Laufeld 1 974a, b*. Samp/esljawed annelids: 7 1 -96LJ, about 3 m below the floar of the quarry, Kettnerites (K) cf. pofonensis, annelid jaws.

GAN NES 3 . Hemse Beds, unit d. Age: Ludlow. Reference: Laufeld 1 974b*; Sundquist 1 9B2c* ; Fredholm 1 988. Sampleljawed annelid: 82- 1 3CB, northernmost part of the bed with large wave marks, no annelid jaws.

GAN NOR 1. Hemse Beds, Hemse Mari uppermost part and Eke Beds, basal part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, b*; Eisenack 1 975, Fig, 30; Laufeld & Jeppsson 1 976; Larsson 1 979; Claesson 1 979; C herns 1 983*; Jeppsson 1 983; Ramskold 1 984, 1 985b; Fredholm 1 988. Samples/jawed annelids: Hemse Beds : 7 1 -26a LJ, loose slab rich i n Dayia navicula (Dayia flags) , representing the uppermost bed in the Hemse Beds, no annelid jaws; 71 -26b LJ, loose slab, (Dayia flags), annelid jaws; 71 -1 21 LJ , about 25 m NW of the reference point, 0.0-0.02 m below the Hemse--Eks boundary, Kettnerites (K) martinssonii, K (K) huberti, annelid jaws; 75-87CB, 0 .30 m below the boundary, no annelid jaws; Eke Beds: 7 1 - 1 22LJ, lowest bed of Eke Beds, no annelid jaws; 75-88CB, the bed immediately above the boundary, no annalid jaws; 7 1 - 1 23LJ , calcarenite poor in fosslis, from the uppermost part of layer 03 in Munthe ( 1 924, Fig 2) and Hede ( 1 925, p. 36 and 38-39. lines 33-38 and 1 - 1 8, respectively), Lanceo/atitesgracilis, Kettnerites (K) bankvaetensis, K (K) huberti, K (K.) polonensis, K. (A) microdentatus, annelid jaws; 7 1 - 1 24LJ, calcarenitic bed i n the calcilutitic layer E i n Mu nthe and Hede op. cit., Kettnerites (K) martinssonii. K (Kl huberti, K. (K) p% nensis, K (A) microdentatus, annelid jaws.

GAN NOR 3 . Hemse Beds, Hemse Mari S E part. Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, b*; Jeppsson 1 983; Fredholm 1 988. Samplesljawed annelids: 7 1 - 1 25U, 90 to 1 00 m SE of the reference point, 0.2 to 0.3 m above the bottom of the ditch. Lanceo/afites gracifis, Kettnerites (K) martinssonii, K. (K.) p% nensis, K (K) huberti, K (A.) cf. sisyphi, annelid jaws; 7 1 - 1 26LJ , uppermost part of the lutitie calcarenite, 0.3 m above the bottom of the small ditch which empties jnto the main drainage ditch, Kettnerites (K.) huberti, K (K.) pOlonensis, K. (A) sisyphi sisyphi, K (A.) microdentatus; annelid jaws.

GARDE 1 . Hemse Beds, unit e . Age: Ludlow, Leintwardinian. Reference:Jeppsson 1 983*; Fredholm 1 988. Samplesljawed anne/ids: 82-34LJ, 1 .64-1 .70 m beJow the reference level , no annelid jaws; 82-35LJ, 0.33-0 . 1 8 m below the reference level, no annelid jaws.


GARORUNGS 1 . Slite Beds, unit f - Rhfpidium tenuistriatum Beds. Age: Early Wenlock. References: Laufeld 1 974a, b"'; Larsson 1 979. Samplesljawedannefids: 73- 1 9LJ, the deepest eroded bed i n the section, fragments; 73-20LJ, 1 .40 m above 73- 1 9 , annelid jaws.

GAROSBY 1 . Hemse Beds, Hemse Mari NW part. Age: Ludlow, Bringewoodian or Leintwardinian. Reference: Jeppsson 1 983*; Fredholm 1 988. Sampfesljawed annelids: B2-27U, very shallow ditch. Lanceolatites gracilis, Kettnerites (A) sisyphi sisyphi, annelid jaws.

GARNUDDEN 1 . Hemse Beds unit a. Age: Ludlow, probabJy Eltonian. References: Laufeld 1 974a, bO; Frykman 1 989. Samp/esljawed annelids: 7547CB, 0.75 m above mean water level and 40 m west of 7546CB, Kettnerites (K.) cf. polonensis, pauli nitid fragments, annelid jaws.

GARNUDDEN 3. 637575 1 67865 ICJ 671 2 7227) ca 471 O m ESE of Anga church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 70 Katthammarsvik.

Low cliff in a small bay at the northern sea shore, ca. 440 m SW of the triangulation point at Garnudden. HemseBeds, unit a. Age: Early Ludlow, possibly Eltonian. Samplesljawed anne/ids: 82-25CB, the second to lowest bed accessible at mean water level, Kettnerites (K.) bankvaetensis, K (K) polonensis, anne lid jaws; 82-26aCB, 0.29-0.32 m above 82-25CB, Kettnerites (K) huberti, K (K.) polonensis, annelid jaws; above former sample, 82-26b CB, ca. 40 mm thick bed with smooth eroded hardground surlace containing some corals, Kettnerites (K) cf. martinssonii, K (K.) huberti, K. (K) p% nensis, annelid

jaws.

GARNUDDEN 4. 637596 1 67937 ICJ6788 7246), ca 5400 m E of Anga church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 70 Katthammarsvik.

Shore exposure at the southernmost tip, 1 25 m N E of the end of the path marked on the topographical map and ca 1 50 m SE of the triangulation point at Garnudden. Hemse Beds, unit a. Age: Early Ludlow, possibly Eltonian. Samplesljawed annefids: 7B· 1 3CB, bed rich in Sphaerorhynchia, Kettnerites (K) bankvaetensis, K (K) martinssonii, K (K) huberti, K. (K.) p% nens/s, anneJid jaws; 78- 1 4CB, bed below 78- 1 3CB, annelid jaws.

G E R ETE 1 . Hemse Beds, Hemse Mari NW part. Age: Ludlow, Bringewoodian or Leintwardinian. Referenees: Laufeld 1 974a, b*; Franzen 1 977; Larsson 1 979; Ramskold 1 983; Jeppsson 1 983; Fredholm 1 988. Samplesljawed annefids: 75-80CB, 20 m east of the bridge and about 1 m above the bottom of the drainage ditch from the lowest of the exposed ( 1 975) beds Lanceofatites gracifis, Kettnerites (A) cf. microdentatus, annelid jaws. 75-81 CB, 0.50 m above 75-80CB, Lanceofatites gracilis, Kettnerites (K) martinssonii, K (A) cf. sisyphi, annelid jaws.

G E R U MSKANALEN 1 . Hemse Bads, Hemse Mari NW part. Age: Ludlow probably Late Bringewoodian, possibly earliest Early Leintwardinian. References: Laufeld 1 974a, b*; LaufeJd & Jeppsson 1 976; Jeppsson 1 982, 1 983*; Fredholm 1 988. Samp/esljawedannelids :7 1 -39LJ, bottom of the ditch, Kettnerites (K) martinssonii. 75-78CB, 2 m W of the bridge, bottom of the ditch, Lanceolatites gracifis, Kettnerites {K) martinssonii, K (K) cf. burgensis, K (A.) sisyphi cf. var. valle, annelid jaws.

GISLE 1 . Hamra Beds, unit a. Age: Ludlow, (Whitcliffian). References: Laufeld 1 974a, b*; Larsson 1 979; Ramskold 1 984. Sampfes/jawedannelids: 76- 1 8CB, uppermost bed, surlace exposure, Kettnerites (K.) bankvaetensis, annelid jaws.

G ISSLAUSE 1 . Slite Beds, unit g . Age: Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979. Samp/esljawed anne/ids: 7?-1 3CB, about 1 .0 m below road surface of the bridge, fragments.

GLASSKAR 1 . Burgsvik Beds, lower part. Age: LudlOw, Whitcliffian. Referenees: Laufeld 1 974a, B*; Larsson 1 979; Jeppsson 1 983. Samp/esljawedannelids: 72-1 BLJ, the about 50 mm thick topmost limestone bed, Kettnerites (K) pofonensis, annelid jaws; 82-1 5CB, grev calcarenite, 0 . 1 4-0.20 m below ground level , Kettnerites (K.) cf. martinssonii, K (K.) p% nensis, K (A) microdentatus, Hindenites naerensis, annelid jaws; 82- 1 6CB, 0.08-0 . 1 4 m below topmost l i mestone bed, soft calcilutite with calcarenitic nodules, Kettnerites (K) p% nensis, annelid jaws, 82-1 7CB, top bed 20-50 mm thick, Kettnerites (K.) cf. martinssonii, K (K) poJonensis including two other varieties, viz. a finely and a coarsely denticulated wide type, Hindenites naerensis, anne lid jaws; 83-1 0LJ, topmost bed, Kettnerites (K) martinssonii, K (K) polonensis and the variety with fine denticles, Hindenites naerensis, annelid jaws.

GLASSKAR 2. Burgsvlk Beds, lower part. Age: Ludlow, Whitcliffian. References: Laufeld 1 974a, bO. SampJesljawed annelids: 72- 1 9LJ, Ihe aboul 50 m m thick topmost l imestone bed, Kettnerites (K) pofonensis, annelid jaws.

GLASSKAR 3. Eke Beds and Burgsviks Beds. Age: Ludlow, Whitcliffian. References: Laufeld 1 974a, b*. Samplesljawedannelids: Eke Beds: 72-20LJ, 0-50 mm above the Eke-Burgsvik boundary, Kettnerites (K.) pofonensis, annelid jaws; 82-20CB, 0 . 1 0-0.07 m below ground level, Kettnerites (K) polonensis, annelid jaws; B2-1 9CB, the about 30 mm thick soft calcilutite, separating the two competent beds 82-20 and 1 B, Kettnerites (K.)

pofonensis, annelid jaws; 82- 1 8CB, the about 40 mm thick, uppermost bed, Kettnerites (K.) cf. martinssonii, K (K) polonensis, K (A) microdentatus, annelid jaws.

GLASSKAR 4. 634905 1 6751 0 ICJ 6 1 54 4596), ca. 1 1 960 m EESE of Burs church. Topographical map sheet 5 1 Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 56 Ronehamn.

Shore exposure on the easternmost point at Glasskår, ca. 2200 m N N E of the house adjacent to the l ight house at Når. Stratified, crinoid-rich limestone with isolated patches of stromatoporoids. Burgsvik Beds. Age: Late Ludlow. Samplesljawed annelids: B3- 1 3CB, the uppermost bed containing stromatoporoids, no annelid jaws; 83- 1 4CB, the uppermost bed 10 m ENE of 83-1 3CB, coarse calcarenite, rich in crinoidal fragments, no annelid jaws.

GLAVES 1 . Hemse Beds, Hemse Mari S E part. Age: Ludlow, Leintwardinian. Referenees: Laufeld 1 974b* ; Jeppsson 1 974:7, 1 983; Larsson 1 979; Fredholm 1 988. Samplesljawed annefids: 75-30LJ, low exposure in the ditch, Kettnerites (K.) martinssonii, K (K.) cf. polonensis, annelid jaws.

G N I SvARD 1 . Upper Visby Beds. Age: Early Wenlock. References: Laufeld 1 974a, bO, Larsson 1 979. Sampfesljawed annelids: 67 -27LJ, loose slab from the excavation of the harbour, fragments; 75-92CB , loose slabs from the excavation of the harbour, Lanceofatites gracilis, Kettnerites (K) martinssonii, K (K) abraham abraham, K (A) sisyphi, annelid jaws.

G N ISvARD 2 . 637763 1 63833 ICJ 2707 7724), ca 3950 m SW of Tofta church. Topographical map sheet 6 I Visby NO. Geological map sheet Aa 1 60 Kl intehamn.

Shore exposure i n the innermost part of the harbour, accessible at low water. Upper Visby Beds. Age: Early Wenlock. Sampleljawed annelids: 79-45LJ, shallow shore exposure, Lanceo/atitesgracilis, Kettnerites (K) martinssonii, K (K) abraham abraham, annelid jaws.

GOORINGS 1 . Halla and Klinteberg Bads, unit a. Age: Late Wen-lock, probably Homerian. References: Laufeld 1 974a, b*; Frykman 1 989. Samplesljawed annelids: Halla Beds: 7 1 -86LJ, Kettnerites (K.) polonensis, K. (A) sisyphi, Hindenites sp. , annelid jaws; Klinteberg Beds: 7?-8CB, loose slab from the ditch excavation, no annelid jaws.

GOORINGS 2. Halla Beds, uppermost part. Age: Late Wenlock, probably Homerian. References: Laufeld 1 974a, b*; Larsson 1 979; Frykman 1 989. Sampfesljawed annelids: 71 -85LJ, Kettnerites (K) cf. polonensis, fragments.

GOGS 1 . Hemse Beds, Hemse Mari S E part. Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, b*; Jeppsson 1 972, 1 974, 1 976, 1 982*, 1 983; Janvier 1 97B; Larsson 1 979; Brood 1 982; Fredholm 1 988. Samp/esljawed annelids: 75-64CB, 0.20 m below the uppermost bed, Lanceolatites gracilis, Kettnerites (K) martinssonii, K (K) huberti, K. (K.) burgensis, annelid jaw fragments; 7 1 -23LJ, excavated material, no annelid jaws.

GOTH EMSHAMMAR 1 . Halla Beds, u n it c, and Klinteberg Beds, unit a. Age: Late Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979; Jeppsson 1 983; Frykman 1 989. Samp/eljawed annelids: 75-36CB, middle part of the section, the most protl1Jding, competent, calcarenitic bed , about 0.5 m above the base of the vertical section, Kettnerites (K.) martinssonii, K (K) polonensis, Hindenites sp., annelid jaws.

GOTH EMSHAMMAR 2. Halla Beds, unit c, and Klinteberg Beds, unit a. Age: Late Wenlock. References: Laufeld 1 974a, bO; Claesson 1 979; Larsson 1 979; Jeppsson 1 983: Frykman 1 989. Samp/es/jawed anne/ids: Halla Beds, unit c : 7 1 -60LJ, the lowest exposed bed in the section, at water level, Kettnerites (K.) cf. bankvaetensis, K (A) microdentatus, annelid jaws; 71 -61 LJ, 1 m above 7 1 -60, Kettnerites (K.) bankvaetensis, K. (K.) martinssonii, K (K) polonensis, annelid jaws; 75-35CB, 0.30 m below the HallaKlinteberg boundary, Kettnerites (K) bankvaetensis, annelid jaws. Klinteberg Beds, un it a : 75-34CB, 0.60 m above the Hal1a-Klinteberg boundary, Kettnerites (K.) bankvaetensis, annelid jaws.

GOTH EMSHAMMAR 3. Halla Beds, unit c, and Klinteberg Beds, unit a. Age: Late Wenlock. References: Laufeld 1 974b* ; Claesson 1 979; Jeppsson 1 983; Frykman 1 989. Samp/esljawedanne/ids: Halla Beds, unit c : 73-23LJ, 1 .80-1 .85 m below the reference level, Kettnerites (K) bankvaetensis, K (K.) cf. p% nensis, annelid jaws; 73-24LJ, 0.80-0.85 m below the reference level, Kettnerites (K) bankvaetensis, other unidentified paulinitid taxa, annelid jaws 73-25LJ, 0.00-0.04 m below the reference level, Kettnerites (K) bankvaetensis, annelid jaws. Kfinteberg Beds, unit a: 73-26LJ, about 10 mm thick and same 1 0-20 mm above, but on the uppermost parts of, the undulati ng, corroded reference level, Kettnerites (K) bankvaetensis, K. (K.) cf. martinssonii, annelid jaws; 73-27LJ, 0 . 1 5--0.25 m above the reference level, representing the uppermost l imestone bed i n the lowest marly beds, Kettnerites (K) martinssonii, K. (A) sisyphi sisyphi, annelid jaws; 73-2BLJ, 0.80-0.85 m above the reference level , Kettnerites (K) bankvaetensis, K (K) martinssonii, K. (A) sisyphi sisyphi, annefid jaws; 73-29LJ, 1 .65 m above the reference level and about the topmost part of the upper marly beds (Hede 1 928)


Kettnerites (K.) maninssonii, annelid jaws; 73·30LJ, 1 .65 m above the reference level, Kettnerites (K.) bankvaetensis, K. (K.l martinssonii, annelid jaws.

GOTHEMSHAMMAR 7 . Halla Beds, unit c. Age: Late Wenlock. References: Laufeld 1 974a, bO; Laufeld & Jeppsson 1 976. Samplesljawed annelids: 77-45PSSFG, Kettnerites (K.l martinssonii, Keltnerites (K.) polonensis var. gandarve, K. (A.) sisyphi sisyphi, annelid jaws.

GOTHEMSHAMMAA 8. Halla Bads, unit c. Age: Late Wenlock. Referenees: Laufeld 1 974a, b' Samplesljawed annelids: 75-37CB, approximately 1 . 75 m below the Halla- Klinteberg boundary, Kettnerites (K.l pa/anensis, K. (A.) sisyphi, annelid jaws.

GAOGAA NSHUVUD 1 . Hamse Beds, unit c. Age: Early Ludlow. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Claesson 1 979; Laufeld & Martinsson 1 98 1 ; Jeppsson 1 982, 1 983; Sundquist 1 982b; Fredholm 1 988. Samplesljawedannelids: 75-51 CB, at water level about 0.3-0.4 m above the bed rich in oriented orthocones, Kettnerites (K) huberti, K. (K.) polonensis, annelid jaws; The samples from 1 98 1 , 8 1 -40LJ-8 1 -43LJ, represent a sequence of about 1 m accessible at low water, with 40 as the lowest and 43 as the highest sample: 8 1 -40LJ, from an anticlinal structure beyond the shore, Kettnerites (K.) huberti, K. (K) polonensis, annelid jaws; 8 1 -41 LJ, Lanceolalites graci/is, Kettnerites (K.) cf. martinssonii, K (K) huberti, K (K) polonensis, annelid jaws; 8 1 -42LJ, Kettnerites (K.) huberti, annelid jaws; 8 1 -43LJ, Lanceofatites graeilis, Kettnerites (K) huberti, K (K) polonensis, annelid jaws.

GRONDALEN l . Klinteberg Beds, lower part. Age: Wen lock-Ludlow. References: Laufeld 1 974a, b* Samplesljawed annelids: 7 1 -51 LJ, about half-way down from the flat upper surface of the island, sample from a 1 0 m thick, thin-bedded u n it, about 1 .5-1 .6 m above its lower contact (a unit with up to 1 m thick beds), Kettnerites (K) polonensis, K. (A) sisyphisisyphi, annelid jaws.

GRUNDARD 2 . 633523 1 64558 IGJ 3 1 07 3444), ca 3420 m WNW of Nas church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa l 52 Burgsvik.

Shore exposure 40 m south of the road and 40 m east of the easternmost part of the small bay. Hemse Beds, Hemse Mari, NW part. Age: Ludlow. Samp/esljawed annelids: 7 1 - 1 46LJ, shallow shore exposure, Kettnerites (K) huberti, K (A) sisyphi sisyphi, annelid jaws.

GRYMUNGS l . Klinteberg Beds, lower-middle part. Age: Late Wenlock. Reference:Jeppsson 1 983* ; Frykman 1 989. Samplesljawed annelids: 79-1 LJ, bottom of the road-side ditch exposure, Kettnerites (K) cf. martinssonii, K (A) sisyphi sisyphi, annelid jaws.

GRYNGE 1 . Hemse Beds, unit d (e?). Age: Ludlow, Leintwardinian. Reference: Fredholm 1 988*. Samples/annelidjaws: 82- 1 6LJ , 5 m W of the navigation marK and about 0.5 m below the contact between the stromatoporoid l imestone and the overlying, dominantly crinoidal limestone, no jawed annelids; 82- 1 7LJ, 5 m S of the navigation mark, from above the contact between the two lithologies, annelid jaws.

GUSTAVSVIK , . Lower Visby Beds, unit b. Age: Late LIandovery. References: Laufeld 1 974b" ; Larsson 1 979. Sampfesljawedannelids: 67-1 LJ, at water level, K (K.) cf. versabiIis, Kennerites (A.) siaelsoeensis, K. (K) abraham abraham, annelid jaws; 79-20LJ, at water level , Kettnerites (A) siaelsoeensis, K (K.) abraham abraham, annelid jaws.

GUSTAVSVI K 2. Lower Visby Beds, unit b(?). Age: Late Liandovery. References: Laufeld 1 974b*. Samplesljawed annelids: 75-1 02CB, uppermost bed, Kettnerites (K) martinssonii, K (A) siaelsoeensis, K (K) abraham abraham, annelid jaws.

GUSTAVSVI K 3. Lower Visby Beds, unit b. Age: Late LIandovery. Reference: Laufeld 1 974b* . Samplesljawed annefids: 79-21 LJ, low section on the beach, Kettnerites (A.) siaelsoeensis, K. (K) abraham abraham, annetid jaws.

GUTENVIKS , . Hemse Beds, unit (c or) d . Age: Ludlow. Referenees: Laufeld 1 974a, bO; Laufeld & Jeppsson 1 976; Fredholm 1 988. Samplesljawed annefids: 7 1 -97LJ, calcilutite, 0.50 m above the bottom of the ditch , Kettnerites (K.) bankvaetensis, K. (K.) martinssonii, K (K) burgensis, annelid jaws.

GUTEVAGEN 2. Hogkl int Beds or Tofta Beds. Age: Early Wenlock. Referenees: Laufeld 1 974b·. Samplesljawed annelids: 69-54LJ, uppermost, stratified bed, rich in algal baUs, annelid fragments.

GYLE 1 . Hemse Beds, unit b. Age: Early Ludlow, Eltonian. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Fredholm 1 988; Frykman 1 989. Samp/esljawed annelids: 75-39CB , 0.25-0.35 m from the base of the section, Kettnerites (K.) cf. martinssonii, annelid jaws; 75-4OC B , 0.40-0.50 m from base of the section, Kettnerites (K) martinssonii, K (K.) polonensis, annelid jaws.

GYLE 2. 636964 1 66890 IGJ 5694 6696), ca 1 1 00 m WWNW ol Ala church. Topographical map sheet 6 J Roma SV. Geological map sheet Aa 1 70 Katthammarsvik.

Ditch section N and S of the intersection of the path and the ditch . Gyle 2 is located just W of the bridge on the NE side of the ditch . Reference level: The boundary between the bluish to brownish, l ight grey limestone and the l ight brownish to l ight grey l i mestone (Hede 1 929:29); the boundary is marked by a surface with corroded pits 1 0-20 mm deep and 30-40 m m wide. Hemse Beds, unit b(?). Age: Early Ludlow. References: Hede 1 929:29-30 l ines 23-31 and 1 -6. Sampleljawedannelids:71 -208LJ, base of the section, 0.8-1 .05 m below the reference level, Kettnerites (K) huberti, K (Kl polonensis, anne lid jaws; 71 -21 1 LJ, 0.5 m above the reference level and 5 m E of the bridge, Kettneriles (K.) martinssonii, annelid jaws.

HAFTINGSKLINT 1 . Upper Visby Beds and Hogklint Beds, unit a. Age: Early Wenlock. References: Laufeld 1 974b", Larsson 1 979. Samplesljawed annefids: Upper Visby Beds: 76-SCB about 1 .0 m a.s . l . , Kettnerites (K) abraham abraham, annelid jaws; 76-9CB about 5 m a.s.I. , Lanceolatitesgracilis var. visby, Kettnerites (K) cf. martinssonii, K (A) sisyphi sisyphi, K (K.) abraham abraham, annelid jaws; 76-1 OCB, approximately the same level as 76-9CB but about l 00 m north of the bioherm, K (K) versabilis, Kettnerites (A) sisyphi sisyphi, K (K) abraham abraham, annelid jaws; 80-3CB, 0.75-0.78 m below the reference level , Kettnerites (K) martinssonii, K (K) abraham abraham, annelid jaws.

HAFTINGSKLINT 4. Lower Visby Beds, Upper Visby Beds, and Hogklint Beds, unit a. Age: Late Llandovery to Early Wenlock. Reference: Hede ( 1 933; 1 to 1 . 5 km S of Haftingsklint). Sampfes/jawed anneJids: Lower Visby Beds, unit et?), or Upper Visby Beds: 84-38LJ, 0 .90 m below sea leve I , annelid jaws; 84-42LJ about 4.25 m a.s. I . , Lanceofatites grad/is, Kettneriles (K) martinssonii, K. (A) microdentatus, K (K) abraham abraham, annelid jaws.

HAGANAS , . Slite Beds, Mari and unit 9 (Ryssnås Limestone). Age: Early Wenlock. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Bergman 1 98 1 , 1 984·; Sundquist 1 98 1 , 1 982a·; Ramskold 1 983; Jeppsson 1 983. Samples/jawed annelids: Slite Mari: ?l-28CB, 4.5 m below reference level Kettnerites (K) martinssonii, K (K.) cf. jacobi, K. (A) sisyphi sisyphi, annelid jaws; 69- 1 0LJ, 1 0-20 mm thick bed at water level, no annelid jaws; 69- 1 1 LJ, immediately above 69- 1 0, Kettnerites (K) cf . bankvaetensis, K (A.) sisyphivar. val le , annelid jaws; Slite Beds, u nit g : 77-29CB, 0.1 m above the reference level (boundary between the Slite Mari and the Ryssnås Limestone), no annelid jaws; 69- 1 3 LJ,

', .0 m below the top of the section, Kenner

ites (A) sisyphi, annelid jaws.

HAGUR 1 . Mulde Beds. Age: Late Wenlock. Reference: Larsson 1 979*. Samples/jawed anne/ids: 8 1 -59LJ, shallow ditch section, KeNnerites (K) martinssonii var. mulde, K. (A) sisyphi sisyphi, annelid jaws.

HAGV I D E 1 . Hemse Beds, Hemse Mari, S E part. Age: Ludlow, Leintwardinian. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 983*; Fredholm 1 9S8. Samples/jawed annefids: 7 1 - 1 29LJ, loose bau Ider, Lanceolatites graci/is, annelid jaws.

HALLAGRUND , . Hogklint Beds, unit c, and Slite Beds, unit c. Age: Early Wenlock. Referenees: Laufeld 1 974a, bO; Jeppsson 1 983. Samples/jawed annelids: Hogklint Beds: 77- 1 9CB, close to the boundary, no annelid jaws. Slite Beds: 79-208LJ, uppermost bed on the southeastern side of the Hållagrundet point, Kettnerites (K) martinssonii, Hindenites gladiatus, annel id jaws. Sl i te Beds: 77-1 8CB, 0.20-0.30 m above the boundary with the Hogklint Beds, no annelid jaws.

HALLBJANNE ,. Eke Beds, lower part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, b" Samples/jawedannelids: 72- 1 7LJ, 0 . 1 -0.2 m below ground level, fragments.

HALLBRO SLOTT 6. Slite Beds, unit c, e , and f. Age: Early Wenlock. Referenees: Jeppsson 1 983* Samples/jawedannelids: Slite Beds, unit c: 73-1 2LJ, 2.55-2.45 m below the reference level, no annelid jaws; Slite Beds, unit e : 73-1 3LJ, 0.05-0 . 1 5 m above t h e reference level, no annelid jaws; Slite Beds, unitf or g(?) : 73- 1 4LJ, 0.80-0.90 m above the reference level, no annelid jaws; 73- 1 5LJ, 2.50 m above the reference level, smaU fragments.

HALUNGE 1 . Klinteberg Beds, unit b. Age: Late Wenlock. References: Laufeld 1 974a, bO; Frykman 1 989. Samples/jawedannelids: 7 1 -93LJ, top of the section, one annelid jaw.

HALU N G E 2. Klinteberg Beds, unit a. Age: Late Wenlock. Referenees: Laufeld 1 974a, b"; Jeppsson 1 983; Aldridge & Jeppsson 1 984; Frykman 1 989. Samples/jawedannelids: 7 1 -94LJ , ditch section, fragments.

HALLSARVE , . Hemse Beds, Hemse Mari , uppermost part, and Eke Beds, lowest part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO; Jeppsson 1 974, p. 1 0, 1 983; Larsson 1 979; Cherns 1 982", 1 983*; Fredholm 1 988. Samples/jawed annelids: 69-26LJ, 0.25-0 . 1 5 m below the reference level, Kettnerites (Kl polonensis, K (K) huberti, annelid jaws;


69-27LJ, 0.05-0. 1 0 m below the reference level and immediately below the bed containing Sha/eria impressa, no annelid jaws; 69-28LJ, bed containing abundant S. impressa, Kettnerites (A) sisyphi klasaardensis, K. (Kl huberti, annelid jaws; 69-29LJ, 0 .3-0.4 m above the reference level, Kettnerites (Kl huberti, other paulinitid taxon, annelid jaws.

HALLS HUK , . Uppar Visby Beds and Hogklint Beds, unit a. Age: Early Wenlock. Referenees: Laufeld 1 974a, b*, Larsson 1 979. Samples/annefid jaws: Upper Visby Beds : n -9CB 1 . 70 m above the reference level, Kettnerites (K.) martinssonii, K (A) sisyphi sisyphi, K (K) abraham abraham, annelid jaws.

HÅLLU DDEN , . 642643 1 68420 IGK 7652 2238), ca 6440 m N N E o f Fleringe church. Topographical m a p sheet 7 J Fårosund SO & NO. Geological map sheet Aa 1 7 1 Kappelshamn.

Fairly low shore exposure, comprising the tip of the land , north of the road. Hogkfint Beds, upper part. Age: Early Wenlock. Sampleljawed annelids: 82-48CB, 0. 1 5 m below the top of the outermost part of the shallow shore exposures, annelid fragments.

HAUGKLI NTAR 2. Klinteberg Beds, lower part. Age:Late Wenlock. References: Laufeld 1 974a, bO; Jeppsson 1 983; Frykman 1 989. Samplesljawed annelids: 70-31 LJ, bedded calcarenite, paul in itid Jaw.

HERRVIK 2. 637048 1 68678 IGJ 7483 6642 ) , ca 3350 m E ol Ostergarn church. Topographical map sheet 6 J Roma 50. Geological map sheet Aa 1 70 Katthammarsvik.

During the excavation of the midd le part of the eastem inner harbour basin i n 1 983, the water was pumped out and the rock sample was taken in the section just outside the newly constructed concrete quay. Hemse Beds, unit d(?). Age: Ludlow. Sampleljawedannelids:83-2CB, about 3 m below sea level, Bluish calcilutite r ich in stromatoporoids with low rel ief, Kettnerites (K.) polonensis, K. (A) cf. fjaelensis, Hindenites sp., annelid jaws.

H I DE 1 . Slite Beds, Slite Mari & Slite g. Age: Wenlock. Referenees: Laufeld 1 974a, bO; Larsson 1 979. Samplesljawedannelids: 73-2LJ, the lowest accessible bed, 1 .95-2 m below the reference level, Hindenitesgladiatus, Kettnerites (K) martinssonii, K (K) huberti, K (A) microdentatus, K. (A) sisyphi sisyphi, Gotlandites slitensis, annelid jaws; 73-3LJ, 1 .37-1 .42 m below the reference level , Kettnerites (Kl martinssonii, Gotlandites slitensis, annelid jaws; 73-4LJ, 0.0-0 . 1 5 m below the reference level, Kettnerites (K) cf. bankvaetensis, K (K) p% nensis, Gotlandites slitensis, annelid jaws. 73-5LJ, 0 .0-0.05 m above the reference level , Gotlandites slitensis, annelid jaws; 73-7LJ, 1 . 1 - 1 .6 m below the top of the section and about 4.5 m above the base of the section, no annel id jaws; n-1 2CB, about 1 m above the base of the section, no annelid jaws; 77-35PSSFG, pau linitid fragments, annelid jaws.

H t D E FISKELAGE 1. Slite Beds, Slite Mari, top and unit g. Age: Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979. Samplesljawed annelids: Slite Mari : 77- 1 4CB, 1 . 1 m below the reference level, Gotfandites slitensis, Kettnerites (K) martinssonii, 77- 1 5CB, about 0.30 m above the reference level, coarse fragmented limestone, no annelid jaws.

HOBURGEN 2. Burgsviks Beds, top part, and Hamra Beds, unit a. Age: Ludlow, WhitcliHian. Referenees: Laufeld 1 974a, b*; Jeppsson 1 974; Claesson 1 979; Larsson 1 979; Laufeld & Martinsson 1 98 1 ; Jeppsson 1 982, 1 983. Samplesljawed annelids: Burgsvik Beds: 69-39LJ, 1 .20-1 .05 m below the reference level, calcarenite (oolite) within the sandstone, annelid jaw; 69-40LJ, 0.SQ-{).60 m below the reference level, Kettnerites (K) d. martinssonii, K (K) huberti, annelid jaws; 69-41 LJ, calcarenite (oolite), 0.35-0.0 m below the reference level, no annelid jaws; Hamra Beds: 69-43LJ, 2.25-2.40 m above the reference level, Kettnerites (K.) polonensis, annelid jaws.

HOBURGEN 3. Burgsvik Beds, Hamra Beds, unit b, c, and Sund re Beds. Age: Late Ludlow. Referenees: Laufeld l 974a, bO; Larsson 1 979. Samples/jawed annelids: Hamra Beds: 7 1 -21 LJ , sample from the roof of the lowest cave, paul initid jaws, annelid jaws.

HOLM HALLAA 1 . 5undre Beds, middle upper part. Age: Latest Ludlow. References: Laufeld 1 974a, b·; Larsson 1 979; Ramskåld 1 983. Samples/jawedannelids: 75-27LJ, crinoidal calcirudite, no annelid jaws; MS905AM, fissure fi l l ing of green and greenish-grey calcilutite, Kettnerites (K.) polonensis, Langeites g/aber, annelid jaws; 75-28LJ, fissure fil t ing, no annelid jaws; 75-29LJ, fissure f i I I ing, roughly Manten's ( 1 971 ) point 1 85 , no annelid jaws.

HOLMHÅLLAR 2 . 631 472 1 651 1 7 IGJ 3507 1 355), ca 5300 m SE of Vamli ngbo church. Topographical map sheet 5 I Hoburgen 50 & 5 J Hemse SV. Geological map sheet Aa 1 52 Burgsvik.

Temporary well excavation, in the small wood just SSE of the Holmhållar boarding-house, 5 m west of the gate across the smatl road. Sundre Beds. Age: Latest Ludlow. Samplesljawedanne/ids: 84-1 1 1 CB, excavated slabs of calcilutite, Lanceolatites sp., f:agments.

HORSNE 3. Halla Bads, unit b. Age: Late(?) Wenlock. Reference: Laufeld 1 974b*; Odin et al . 1 986. Samplesljawed annelids: 75-36KL, annelid jaws; 82-8CB, the uppermost 0 .05-0.07 m of the section, immediately above the small bioherm, no annelid jaws; 82-9CB , about 4 m E of 82-8 i n a calcilutitic pocket within the middle part of the reet, about 0.7 m


above the water in t h e drainage ditch, annelid jaws; 82-1 acs. dose lo 82-9 i n the same pocket, Kettnerites (K.l martinssonii, annelid jaws; 82- 1 1 CB, very soft calcilutite in the same pocket, annelid jaws; 82- 1 2CB, competent calcitutite, Kettnerites (K.) martinssonii, annelid jaws.

HORS N E 5. Halla Beds, unit b. Age:Late(?) Wenlock. References: Laufeld 1 974a, b*, Samples/jawed annelids: 75-52CB, excavated material, Kettnerites {K.l martinssonii, K. (K.l aH. jacobi, K. (A) sisyphi sisyphi, anne lid jaws; 75-53CB, excavated material, Kettnerites (K.l cf. martinssonii, K. (K.l aff. abraham isaae, annelid jaws; 75-55CB, excavated material, annelid jaws.

HORSNE 6. Halla Beds, unit b. Age:Late{?) Wenlock. Reference: Larsson 1 979; Ramskold 1 985b. Samples/jawed annelids: 74·37KL, annelid jaws; 74·39KL, Kettnerites sp. , annelid jaws; 75-54KL, annelid jaws.

H U M M E LBOSHOLM 1 . Eke Beds, lower part. Age: Ludlow. References: Laufeld 1 974a, b·. Sampleljawed annefids: 76·1 1 CB, shaliow ditch section, Lanceolatites gracms, Kettnerites (K) martinssonii, K (Kl huberti, annelid jaws.

HUNN INGE 1. Klinteberg Beds, lower-middle part. Age: Late Wenlock. References: Laufeld 1 974a, b'; Larsson 1 979; Jeppsson 1 983; Frykman 1 988. Samplesljawedannelids:71 -3LJ, about 2.5-3 m above the base of the section, no annelid jaws.

HUSRYGGEN 3. Hamra Beds, unit b. Age: Ludlow, Whitcliffian. Reference: Larsson 1 979. Samplesljawedannefids: 7 1 - 1 66LJ , lowest exposed bed, Lanceofatites sp. , annelid jaws; 7 1 - 1 68LJ, 4.4 m above 7 1 - 1 66 , paul initid fragment, annelid jaws.

IREVIKEN 1 . Lower Visby, Upper Visby, and Hogklint Beds, unit a. Age: Late Llandovery to Early Wenlock, References: Laufeld 1974a, b"; Ramskold 1 983, 1 984. Samplesljawedannelids: Lower Visby Beds, unit e : 76-4GB, 0.5 m below the reference leve l , K. (K) versabilis, K. (K) abraham abraham, annelid jaws; Hogklint Beds, unit a: 76-5GB, loose slab from the bedded, l ight brown calcarenite between the reef bodies, Kettnerites (K.) abraham cf. abraham, annelid jaws.

IREVIKEN 2. Lower Visby Beds, Upper Visby Beds, and Hogklint Beds, unit a. Age: Late Llandovery{?) to Early Wenlock. References: Laufeld 1 974b", Larsson 1 979. Sampfes/jawedannelids: Lower Visby Beds: 76-2GB, about 0.3 m above sea level, KeNnerites (K.) abraham abraham, annelid jaws; Upper Visby Beds: 76-3CB about 6 m below the reference level, Kettnerites (K,) martinssonii, K. (Kl abraham abraham, annelid jaws.

IREVIKEN 3. Lower Visby Beds, Upper Visby Beds, and Hogklint Beds. Age: Late Llandovery to Early Wenlock. Referenees: Laufeld 1 974a, b'; Larsson 1 979; Odin et al . 1 984", 1 986. Samplesljawedannefids: Lower Visby Beds unit b : 79� 1 5LJ, about 4 m below the reference level , Kettnerites (K) versabifis, K. (A.) siae/soeensis, K (K) abraham abraham, annelid jaws; 79- 1 6LJ, 0 . 1 -0.2 m below the reference level , Kettnerites (K) abraham abraham, annelid jaws; 82-2LJ, 0.02-0 . 1 5 m above the reference level, Kettnerites (A.) siaelsoeensis, K (K) abraham abraham, annelid jaws; 82-3LJ, 0.85-0.90 m a.s . I . , Kettnerites (A.) siae/soeensis, annelid jaws; Lower Visby Beds unit c: 81 -6LJ, 2.36-2.46 m above the reference level , Kettnerites (K.) cf. versabilis, Lanceolatites cf. graeilis, Kettnerites (K.) abraham abraham, annelid jaws; 82-5LJ, 1 .9 1 -2.06 m above the reference level, no annelid jaws; 82·6LJ, 2 . 1 6-2.26 m above the reference level, Kettnerites (K.) martinssonii, K (K.) abraham abraham, annelid jaws; 82·8LJ, 2 . 9 1 -3.01 m above reference level, Lanceolatites gracilis, Kettnerites (K) martinssonii, K abraham abraham, annelid jaws; Lower Visby Beds unit d : 81 -7LJ. 3.96 m above the reference level , Kettnerites (K.) versabilis, K. (K.) martinssonii, K. (K.) abraham abraham, annelid jaws; Lower Visby Beds unit e: 82-9LJ, 4 .21-4.26 m above the reference level, fragments; 82-1 OLJ, 5 . 1 -5.3 m above the reference level, KeNnerites (K) abraham abraham, annelid jaws; 83-25LJ, 5.32-5.44 m above the reference level, Kettnerites (K) abraham abraham, annelid jaws; Upper Visby Beds: 79-1 8LJ, 9.25 m above the reference level, Lanceolatites gracifis, Kettnerites (K.) abraham abraham, annelid jaws; 82-1 1 LJ , either 5.5 or 6.2 m above the reference level, Lanceo/atites graci/is, Kettnerites (K) martinssonii, K (K.) abraham abraham, annelid jaws.

JAKOBSBERG 1 . Sl ite Beds, unit g . Age: Wenlock. References: Laufeld 1 974a, b'; Larsson 1 979. Samplesljawedannelids: 72-4LJ, from the lower part of the eroded bed , same level as 72-47SL (Laufeld 1 974a), 0 .1 0-0.30 m below the 'step' (along the path) in the section, argillaceous calcilutite, fragments.

JUVES 1 . Hamra Bads, unit c, and Su ndre Beds, lower part. Age: Latest Ludlow. References: Laufeld 1 974b*; Larsson 1 979. Samples/jawed annelids: Hamra Beds: 76·25GB, s l ightly more than 2 m above the base of the section, anne lid jaws.

JUVES 2. Hamra Beds, unit c, and Sundre Beds, lower part. Age: Latest Ludlow. Referenees: Laufeld 1 974a, b'; Larsson 1 979; Jeppsson 1 982*, 1 983.


Samples/jawed annelids: Hamra Beds: 7 1 -20LJ , 5 . 1 7-5, 1 2 m below the boundary between the Hamra and Sundre Beds, pau l init id fragment, fragments; 69-44U, 5 . 0 1 -4.75 m below the boundary between the Hamra and Sundre Beds, fragments; 69�46LJ, 3.53-3.43 m below the boundary between the Hamra and Sundre Beds, fragments.

The stratigraphic levels of LJ samples from the Juves localities 2, 3, and 5 refer to the boundary between the Hamra and Sundre Beds which does not agree with the reference level at Juves 2 designated by Laufeld ( 1 974b). For further information, see (Jeppsson 1 982, p. 1 7, Fig. 2, the composite section for Juves with the levels in italics).

JUVES 3. Hamra Beds, unit c, and Sundre Beds, lower part. Age: Latest Ludlow. References: Laufeld 1 974a, bO; Larsson 1 979; Glaesson 1 979; Jeppsson 1 982', 1 983. Sampfes/jawedannelids: Hamra Beds: 77-3LJ, 2.38-2.23 m below the reference level , no annelid jaws; 77- 1 9LJ, 1 .55-1 .50 m below the reference level, annelid jaws; 69-47LJ, 1 .03-0.90 m below the reference level, Kettnerites (K) huberti the slender type, annelid jaws; 69-49LJ, 0.0-0 . 1 5 m below the reference level, pau l in itid fragments, annelid jaws; Sundre Beds : 69-50 U , about 3 m above the reference level, fragments.

JUVES 4. Sundre Beds, lower part. Age: Latest Ludlow. References: Laufeld 1 974a, b'; Larsson 1 979; Jeppsson 1 982', 1 983. Samplesljawedannelids; 76-23GB, Kettnerites {K.) bankvaetensis, paul i nitid sp. indet. , annelid jaws.

JUVES 5. Hamra Beds, unit c. Age: Latest Ludlow. References: Laufeld & Jeppsson 1 976*; Jeppsson 1 982", 1 983. Samp/esljawed annelids: 7 1 - 1 82LJ, 6.57-6.50 m below the reference level, no annelid jaws; 77- 1 1 LJ , 5.52-5.43 m below the reference level, no annelid jaws; 77-1 OLJ, 4.53-4.48 m below the reference level, no annelid jaws.

KAKHUSE 1. 6371 68 1 65477 ICJ 4298 701 2), ca 3060 m NW of G u ldrupe church. Topographical map sheet 6 J Roma SV. Geological Map sheet Aa 1 60 Klintehamn.

Shallow ditch section, on the S E side of the small road from Våte to Vikfau, about 1 75 m NE of the cross-roads at point 41 ,68. Klinteberg Beds, lower part. Age: Latest Wenlock - earliest Ludlow. Sampfesljawed annelids: 78-9CB, 0.25 m below the uppennost surface of the l imestone, no annelid jaws.

KÅLLDAR 1 . Hemse Beds, Hemse Mari NW part. Age: Ludlow. References: Laufeld 1 974a, b·. Sampleljawed annelids: 7 1 -34LJ, excavated material from ditch at Vissne myr, about 3 km south of Linde church, dumped at Kalldar 1 , Lanceolatites cf. gracilis, Kettnerites (A.) sisyphisisyphi, annelid jaws; 75-75CB, excavated material from the same locality as 7 1 -34LJ, KeNnerites (A.) microdentatus, annelid jaws.

KÅLLDAR 2. Hemse Beds, lower-middle part. Age: Ludlow, Bringewoodian or possibly earliest Leintwardinian. References:Jeppsson 1 983'; Fredholm 1 988. Samplesljawed annelids: 7 1 -35LJ, middle part of the section, Kettnerites (A.) sisyphi sisyphi, annelid jaws.

KAMBS 5. 640444 1 65660 ICK 4741 0253), 3770 m WSW of Martebo church. Topographical map sheet 7 J Fårosund SV & NV. Geological map sheet Aa 1 83 Visby & Lummelunda.

Drainage ditch section on the northern side of the ditch, im mediately E of the bridge. Tofta Beds. Age: Early Wenlock. Reference: Hede 1 940, p. 35, lines 3-5 from below. Samples/jawed annelids: 75-28CB, topmost bed in the section, no annelid jaws; 82-42LJ, topmost bed in the section, a fragment.

KAPELLSHAMN 1 : Hogklint Beds unit b. Age: Early Wenlock. Reference: Laufeld 1 974a, bO, Larsson 1 979. Samplesljawed annelids: 77- 1 1 C B, 0.5 m below the top of the section, Kettnerites (K.) martinssonii.

KAPELLUDDEN 1 . 634751 1 67 1 32 (CJ 5764 4472), ca 8500 m ESE of Burs church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 56 Ronehamn.

Shore exposure 200 m N of Kapellet in Kapelluddens fiskelage marked on the topographical map sheet. The exposure is also marked on the geological map sheet. Burgsvik Beds. Age: Ludlow. Referenees: Hede 1 925a, p. 3 1 . Sampleljawed annelids: 83- 1 6LJ, shallow shore exposure, Kettnerites (K.) polonensis, K (K.) huberti, Hindenites naerensis, annelid jaws.

KÅRNE 3. Hemse Beds, Hemse Mari, uppermost part and Eke Beds, basal part. Age: Ludlow, Late Leintwardinian. l1eference:Jeppsson 1 983"; Fredholm 1 988. Samples/jawed annelids: Hemse Beds: 7 1 - 1 95LJ, ditch exposure, 1 33 m south of the contact between the Hemse and Eke Beds, Kettnerites (K.) burgensis, Langeites glaber, annelid jaws; 7 1 -1 96L.J, 8 m south o f t h e contact, n o annelid jaws, 71 ·1 97LJ , Shaleria impressa-rich level, top Hemse, 0 .01-0.04 m below the contact, Kettnerites (K.) huberti, annelid jaw; Eke Beds: 7 1 - 1 98LJ, 0.0-0.06 m above the contact, Kettnerites (A.) sisyphi klasaard,ensis, K. (K) huberti, annelid jaws; 7 1 - 1 99LJ, bryozoan·rich conglomerate, 0.06-0 . 1 6 m above the contact, Kettnerites (A.) microdentatus, annelid jaws; 7 1 -200LJ, top of the section, no annelid jaws; 7 1 -202LJ, 5 m N of the road, no annelid jaws; 71 -203LJ, N of Karne, no annnelid jaws.

KATTELVIKEN 5. Hamra Beds, unit b. Age: Ludlow, Late Whitcliffian or slightly younger.

References: Laufeld & Jeppsson 1 976'; Larsson 1 979; Jeppsson 1 983. Samp/esljawed annelids: 7 1 - 1 78LJ, 3 m below the top ot the section, Kettnerites (K.) polonensis, paulinitid sp. indet., annelid jaws; 7 1 - 1 79LJ, see Laufeld & Jeppsson 1 976, annelid jaws; 77-33LJ, 3 m below top of the section, Lanceolatites gracilis, Kettnerites (K) martinssonii, K (K.) huberti, K (K.) polonensis, Hindenites sp., annelid jaws; 7 1 - 1 76LJ, 1 m below the top of the section, Kettnerites (K.) bankvaetensis, K. (K.) polonensis, a'hnelid jaws; 7 1 - 1 75LJ , uppermost exposed bed, paulinitid fragments, annelid jaws.

KATIHAMMARSVIK 1 . Hemse Beds, unit a. Age: Early Ludlow. References:Laufeld 1 974a, bO; Laufeld & Martinsson 1 98 1 ; Jeppsson 1 983; Fredholm 1 988. Samplesljawed annelids: 67-30LJ, shore exposure, annelid jaws; 75-50CB, upper part of the shore exposure, annelid jaws.

KAUPARVE 1 . Hamra Beds, lower-middle part. Age: Ludlow. References: Laufeld 1 974a, bO. SampJeljawed annelids: 76· 1 6CB, 0.2-0.3 m below ground level , Kettnerites (K.) bankvaetensis, K. (K.) martinssonii, annel id jaws.

KLASARD 1 . Hemse Beds, Hemse Mari SE part. Age: Ludlow, Leintwardinian. References:Jeppsson 1 983"; Fredholm 1 988. Samplesljawedannelids: 7 1 · 1 50LJ, shore exposure, Lanceolatites graci/is, Kettnerites (K) burgensis, K (A.) sisyphi klasaardensis, K (K.) huberti, annelid jaws.

KLINTEBERGET 1 . Klinteberg Beds, lower and middle parts. Age: Late Wenlock. Referenees: Laufeld 1 974a, bO; Claesson 1 979; Larsson 1 979; Laufeld & Martinsson 1 98 1 ; Jeppsson 1 982, 1 983; Ramskold 1 983; Frykman 1 989. Samplesljawedannelids: 67-29LJ, about 8 m below the top of the cliff, Kettnerites (K) polonensis, annel id jaws; 72-23KL, pockets of greenish- and brownish-grey calcilutite, Kettnerites (K) martinsson;;; 75- 1 9GB, about 2 m below the top of the cliff section, annelid jaws; 75-20CB, about 3 m below the top of the cliff section, annelid jaws; 75-21 CB, lutitic calcirudite above the thickly bedded l imestone beds, close to tap of the section, Hindenites angustus, annelid jaws.

KLINTEEN KLAVEN 1 . Slite Beds, Slite Siltstone. Age: Wenlock. Referenees: Laufeld 1 974a, bO; Sivhed 1 976; Bergman 1 979b, 1 980a. Samplesljawed annelids: 75-24GB, 0.4 m from the top of the section, Kennerites (A.) sisyphi cf. var. valle, annelid jaws.

KLI NTHAG EN 1 . Slite Beds, unit g . Age: Early Wenlock. Reference: Jeppsson 1 983�. Samplesljawed annelids: 73- 1 LJ , immediately below the calciruditie bed (reefal detritus), no annelid jaws.

KLUVSTAJN 1 . Slite Beds, unit b. Age: Early Wenlock. References: Laufeld 1 974a, b'; Jeppsson 1 983. Samp/es/jawed anne/ids: 73- 1 6LJ, excavated material from a trench, paul initid fragments.

KLUVSTAJN 2. 638527 1 64352 ICJ 3285 8445), ca 2820 m WSW of Vasterhejde church. Topographical map sheet 6 I Visby NO. Geological map sheet Aa 1 83 Visby & Lummelunda.

Shallow ditch exposure, 6 m NW of the SE end of the truncation of the corner of the tield and about 575 m NNW of point 54,4. Slite Beds, unit a(?). Age: Early Wenlock. Samples/jawed annelids: 73- 1 7 LJ , 0.1 m thick exposure, Kettnerites (K) cf. martinssonii, annelid jaws,

KODINGS 3. Hemse Beds, Hemse Mari S E part. Age: Ludlow. References: L::aufeld 1 974a, b"; Larsson 1 979. Sampfesljawedannelids: 75-82G B, ditch exposure, annelid jaws.

KORP KU NT 1 . Upper Visby Beds and Hogklint Beds, unit a. Age: Early Wenlock. References: Laufeld 1 974a, bO, Larsson 1 979. Samp/esljawed annelids: Upper Visby Beds: 67-2LJ, about 5 m below the reference level, Kettnerites (K) cf. versabifis, Lanceolat;tes sp., Kettnerites (K) martinssonii, K (K.) abraham abraham, annelid jaws; 75-5GB, immediately below the reference level, Kettnerites (K) martinssonii, K (K) abraham abraham; Hogklint Beds, unit a : 75-4GB immediately above the reference level, no annelid jaws.

KRAK FOT 1 . Klinteberg Beds, unit d{?) . Age: Latest Wenlock -earliest Ludlow. Reference: Frykman 1 989". Samples/jawed annelids: 7 1 - 1 53LJ, bottom of the ditch, 1 00 m east of the road, Kettnerites (K) bankvaetensis, annelid jaws.

KRASSE 1 . Klinteberg Beds, middle part. Age: Latest Wenlock -earliest Ludlow. References: Laufeld 1 974a, bO; Frykman 1 989. Samples/jawed annelids: 78-1 OCB, uppermost bed i n the section, no annelid jaws.

KROKEN 1 . Burgsviks Beds and Hamra Beds. Age: Ludlow. References: Laufeld 1 974a, b·. Sampleljawed annelids: Burgsvik Beds: 72-2 1 LJ , 0 . 1 -0 . 05 m below the Burgsvik-Hamra boundary, paul in itid sp., annelid jaws.

KROKEN 2. Burgsvik Beds and Hamra Beds. Age: Ludlow. References: Laufeld 1 974a, b·. Samp/eljawed anne/ids: 72-22LJ, lateral of a Hamra bioherm, Kettnerites (K) bankvaetensis, K. (K) martinssonii, K (K.) huberti, K. (K.) cf. polonensis, K. (A.) microdentatus, annelid jaws.


KROKEN 3. 634784 1 6 7464 (CJ 6098 4480). ca 1 1 580 m ESE of Burs church. Topographical map sheet 5 I Hoburgen NO & 5 J Hamse NV. Geological map sheet Aa 1 60 Kl intehamn.

Shore exposure on the narrow point just east of the 'n' in 'Kroken' on the topographical map sheet. Hamra Bads. Age: Ludlow. Samplesljawed annelids: a3- 1 2GB, shore exposure, no annelid jaws.

KULLANDS 1 . Hamse Beds, Hemse Mari NW part. Age: Ludlow, Bringewoodian or Early Leintwardinian. Referenees: Laufeld 1 974a, b*; Laufeld & Jeppsson 1 976; Larssen 1 979; Jeppsson 1 983. Samplesljawed annelids: 75-79CB, uppermost bed at the botlom of the water hole, covered by about 0 . 1 -0.2 m sail , Kettnerites (K) cf. martinssonii, annel id jaws.

K ULLANDS 2 . 635370 1 65 1 52 (CJ 3838 5237), ca 1 370 m SSW of Gerum church. Topographical map sheet 6 J Roma SV. Geological map sheet Aa 1 64 Hemse.

Ditch section, at the intersection of the private road (not marked on the topographical map sheet) and the ditch, about 1 40 m W of the western most house (on the topographical map sheet) at Kullands. Hemse Beds, Hemse Mari NW part. Age: Ludlow. Reference: Fredholm 1 988. Sample/jawedannelids:84-31 1 OF, Kettnerites(K) bankvaetensis, K. (K.) martinssonii, K (K.) burgensis, K (A) fjaelensis, annelid jaws; 84-31 2DF, Lanceolatites graeilis, Kettnerites (K) martinssonii, K. (A) fjae/ensis, anne lid jaws.

KUPPEN 1 . Hemse !3eds, unit d . Age:Ludlow, Leintwardinian. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 982, 1 983; Kershaw 1 987. Samp/es/jawed annelids: 8 1 -44LJ, 0.0-0. 1 1 m above the reference level, no annelid jaws; 8 1 -45LJ, 0 . 1 1 --0 . 1 4 m above the reference level, no annelid jaws.

KUPPEN 2. Hemse Beds, unit d . Age: Ludlow, Leintwardinian. Referenees: Laufeld 1 974a, bO; Kershaw & Riding 1 978; Laufeld & Martinsson 1 98 1 ; Kershaw 1 98 1 , 1 987; Jeppsson 1 982, 1 983; Fredholm 1 988. Samp/es/jawed anne/ids: 8 1 -46LJ, no annelid jaws; 8 1 -47LJ, no annelid jaws.

KvANNVATEN 1 . Slite Beds, Pentamerus goth/andieus Beds or slightly younger. Age:Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979. Samp/es/jawedannefids: 77- 1 C B, about 2 m below the top of the section, no anne lid jaws.

LAMBSKVIE 1 . Hemse Beds, unit c. Age:Ludlow, probably Bringewoodian. Referenees: Laufeld 1 974a, bO; Larsson 1 979. Samp/esljawed annelids: 74- 1 8KL, Kettnerites (K) p% nensis; 74-62KL, Kettnerites (K) polonensis; 75-42CB, loose slab of excavated material, Kettnerites (K) p% nensis, anne lid jaws; 75-44CB, top bed on the eastern side, Kettnerites (K.) huberti, annelid jaws.

LANG HAM MARS HAMMAR 1 . H6gklint Beds, lower-middle part. Age: Early Wenlock. Referenees: Laufeld 1 974a, bO, Jeppsson 1 983. Samp/es/jawedannelids: 77-26CB , jawed annelids.

LANGHAMMARSVIKEN 2. 643385 1 69795 (CK 9079 2872), ca 7750 m N of Fåra church. Topographical map sheet 7 J Får6sund SO & NO. Geological map sheet Aa 1 80 Fåra.

Shallow exposure with large wave marks at the sea shore NW of the westemmost extension of Langhammarsviken. Hogklint Beds, lower-middle part. Age: Early Wenlock. Referenees: Hede 1 936 : 1 6 l ines 6-9; Bergman 1 979b; Larsson 1 979. Samp/e/jawed annelids: 78-1 7CB, surfaee exposure, sample of the large wave-marked material, Kettnerites (K.) martinssonii, K. (K.) abraham isaac, annel id jaws.

LASSOR 1 . Hemse Beds, upper part. Age: Ludlow. Referenees: Laufeld 1 974 a, bO ; Fredholm 1 988. Samp/esljawedannelids: 72-8LJ, 0.50-0.60 m below the top of the section (same level as sample 72-5 1 S L in Laufeld 1 974a), no annelid jaws; 78-6CB, loose slabs, fragments.

LAU BACKAR 1 . Eke Beds, Rhizophylfum limestone. Age: Late Ludlow. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Ramsk61d 1 983, 1 984, 1 985b.

.

Samples/jawed annefids: 72-27KL, annelid jaws; 77-30CB, above the bottom of the excavation and about 0.7 m below ground level , Kettnerites (A) cf. sisyphi, annelid jaws.

LAUTER 1 . Hagklint Beds, lower middle part. Age: Early Wenlock. Referenees: Laufeld 1 974a, bO. Samp/esljawedannelids: 77-24CB, suriace exposure, the topmost 20--50 mm, Kettnerites (K) martinssonii, K (A) sisyphi sisyphi, K. (K) abraham cf. isaac, annelid jaws.

LAUTERHORNSVIK 2 . Hagklint Beds, lower-middle part. Age: Early Wenlock. '

References: Laufeld 1 974a, bO, Jeppsson 1 983. Samp/esljawedannefids: 73-73LJ, low shore exposure, Kettnerites (K) martinssonii, annelid jaws.

LAUTERHORNSVIK 3. Hagklint Bads, lower-middle part. Age: Early Wenlock. References: Larsson 1 979', Jeppsson 1 983. Samplesljawed annelids: 73-72LJ, shallow ditch exposura, Kettnerites (K.) abraham cf. isaac, annelid jaws.

LERBERGET 1 . Slite Beds, Slite MarI, Lerberget Mari and Penfamerus gothlandicus Beds. Age: Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 983. Samp/es/annefid jaws: Lerberget Mari: 79-37LJ, lowest exposed bed, Kettnerites (A) sisyphi sisyphi, anne lid jaws; 70-37LJ, 0.30 m above the base of the section, fragments; 7D-38LJ, 0 .95 m above the base of the section, Kettnerites (A) sisyphi sisyphi; 70-39LJ, 2.75 m above the base of the section, bad rich in Ketophyffum, fragments; Pentamerusgothlandicus?: 70-40LJ, 5. 1 5 m above the base of the section, pau li nitid sp. indet., fragments; 70-41 LJ, 7.0 m above the base of the section, pau li nitid sp. indet.; 70-42LJ, 9 . 1 5 m above the base of the section and about at the top of the mari, Kettnerites cf. abraham, K (A) cf. sisyphi, annelid jaws; 70-43LJ, about 5 m above the base of the succeeding limestone, no annelid jaws; 7D-44LJ, about 15 m above the base of the l imestone, no annel id jaws.

L lCKERS 1. 641 607 1 66265 (CK 5399 1 335), ca 4840 m N of Stenkyrka church. Topographical map sheet 7 J Fårasund SV & NV. Geological map sheet Aa 1 83 Visby & Lummelunda.

Locality comprising the beach and adjacent cl iff, the area batween the bioherms. Lower Visby Beds, unit b or c, on the shore. Age: Late Uandovery. Samples/jawed annelids: 8 1 - 1 DU, 0.1 m above mean sea level, Lanceolatites cf. gracifis, Kettnerites (K) abraham abraham, annelid jaws.

L1CKERSHAMN 2. Upper Visby Beds and H6gklint Beds, un its a and b(?). Age: Early Wenlock. Reference: Laufeld 1 974a, bO, Larsson 1 979. Samp/es/jawed annelids: Upper Visby Beds: 73-54LJ, about 0.9 above the base of the section, Kettnerites (K) abraham abraham, annelid jaws; 73-55LJ, 2.65 above the base of the section, a 60--80 mm thiek bed, Lanceolatites gracilis, Kettnerites (A) sisyphi sisyphi, annelid jaws; Upper Visby Beds(?); 73-56LJ, 4 .35 m above the base of the section, Kettnerites (K.) martinssonii, K. (K.) abraham abraham, annelid jaws; Hagklint Bads, probably unit a; 73-58LJ, 5 .35 m above the base of the section, a 75 mm thick bad immediately above a bentonite(?), Kettnerites (K.) martinssonii, annelid jaws; 73-59LJ, 8.30 m above the base of the section, Kettnerites (K.) martinssonii, K (A) sisyphi sisyphi, K. (K.) abraham abraham, anne lid jaws; 75- 1 03CB, reef level in lowest Hagklint, no annelid jaws; 75- 1 06CB, about 4 m below the reference level, no annelid jaws; H6gklint Bads, unit b(?); 73-60LJ, 9.85 m above the base of the section, Kettnerites (K) martinssonii, K. (A) sisyphi sisyphi, K (K.) abraham cf. isaae, annelid jaws; 73-61 LJ, 1 1 .90 m above the base of the section, Kettnerites (K) martinssonii, annelid jaws; 73-62LJ, 1 4 . 05 m above the base of the section, Kettnerites (K) martinssonii, annelid jaws.

L 1 K M I D E 2. Hemse Beds, Hemse Mari SE. Age: Ludlow, Late Leintwardinian. Reference: Fredholm 1 988*. Samplesljawed annelids: 82-28LJ, shallow ditch, Lanceo/atites gracilis, Kettnerites (K.) burgensis, K (A) Ifaelensis, annelid jaws. LILLA HALLVARDS 1 . Hemse Beds, Hemse Mari NW part. Age: Early Ludlow. References: Laufeld 1 974a, bO; Larsson 1 979; Jaeger 1 98 1 ; LaufeId & Martinsson 1 98 1 ; Jeppsson 1 983; Fredholm 1 988. Samples/jawed annelids: 71 - 1 43LJ, ditch exposure, Lanceo/atites gracilis, Kettnerites (K.) cf. martinssonii, K. (A) sisyphi, annelid jaws.

LI LLA HALLVARDS 4. Hemse Beds, Hemse Mari NW part. Age: Early Ludlow. References:Larsson 1 979*; Fredholm 1 988. Samples/jawed annefids: 84-3 1 4DF, Lanceo/atites gracilis, Keltnerites (K.) martinssonii, K. (A) microdentatus, anne lid jaws.

L I N D E 1 . Hemse Beds, upper part. Age: Ludlow. References: Laufeld 1 974a, b'; Larsson 1 979; Ramskald 1 983. Sampfes/jawedannelids: 7 1 - 1 1 2 LJ, about 2 m above road suriace, no annelid jaws; 71 - 1 1 4LJ, on the slope about 0.5 m above the edge of the section, no annelid jaws.

L INVIKEN 2. 633493 1 64598 (CJ 3 1 46 341 9), ca 2950 m WNW of Nas church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 52 Burgsvik.

Shore exposure 750 m SSE of the solitary house, Glose to the stone fenee that starts by the cattle grid on the road N E of the locality. Hemse Beds, Hemse Marl NW part. Age: Ludlow. References: Hede 1 9 1 9 , pp. 1 7- 1 9 , Ioc. 6 ; Hede 1 942, p. 20, Ioc. 4C. Samples/jawed annefids: 7 1 - 1 47LJ , shore exposure, Kettnerites (K.) huberti, K. (K) burgensis, annelid jaws.

LJ UGARN 1 . Hemse Beds, upper part, probably unit d . Age: Ludlow. References: Laufeld 1 974a, bO; Watkins 1 975; Jeppsson 1 983; Fred holm 1 988. Samplesljawed annelids: 79-51 LJ , shore exposure, no annelid jaws.

LOGGARVE 1 . Mulde Beds, uppermost part and Klinteberg Beds, lowest part. Age: Late Wenlock. References: Jeppsson 1 982, 1 983*; Frykman 1 989. Samples/jawed annelids: Mulde Bads: 77-22LJ, 0.50 above the reference level. A comparison with Loggarve 2 has shown that only the upper 0 .75 m of the strata balongs to the Klinteberg Bads : Kettnerites (K.) polonensis, K. (A) sisyphi sisyphi, annelid jaws; Klinteberg Bads: 77-23LJ, 2.5 m above the reference level, no ann:lid jaws.

LOGGARVE 2 . Mulde Beds and Klinteberg Beds, lowest part. Age: Late Wenlock.


T h e contact batween t h e basal 0 . 1 m of soft Glay a n d t h e succeeding harder beds remains the best reference level (Lennart Jeppsson, personal commun ication). The contact is found about 2 m below the base of the Klinteberg Bads, defined by the initial occurrenee of the brachiopod Conchidium conchidium. References: Jeppsson 1 982*; Ramsk61d 1 985b; Frykman 1 989. Samples/jawed annefids: Mulde Beds: 82-21 LJ, 1 .98--2.05 m below the reference level, Kettnerites (K.) martinssanii, K (A) sisyphi sisyphi, annelid jaws; 82-22LJ, 1 .80--1 .72 m below the reference level, Kettnerites (K.) martinssonii, K (K.) p% nensis var. gandarve, annelid jaws; 82-23LJ , 0.87--0.80 m below the reference level, Kettnerites (K) sp. , annelid jaws; 82-24LJ, top Mulde, 0 . 1 0-0.00 m below the reference level, Kettnerites (A) sisyphi sisyphi, annelid jaws; 82-25LJ, base of Klinteberg Bads, 0.02--0.05 m above the reference level, Kettnerites (K) martinssoniivar. mulde, Hindenites angustus, annelid jaws.

LUKSE 1 . Hemse Bads, Hemse Mari NW part. Age: Ludlow. Referenees: Laufeld 1 974a, bO ; Larsson 1 979; Ramsk61d 1 983; Jeppsson 1 983; Fredholm 1 988. Samples/jawedannelids :7 1 - 1 2LJ , about 0.2 m below the top of the culvert, Lanceolatitesgracilis, Kettnerites (A) sisyphisisyphi, paulinitid sp., annelid jaws; 7 1 - 1 3U , loose s1abs, Lanceolatitesgracifis, Kettnerites (A) fjae/ensis, annelid jaws.

LYR UNGS 1 . Hemse Beds, probabJy middle part. Age: Ludlow. Referenees: Laufeld 1 974a, bO ; Fredholm 1 988. Samp/esljawed annelids: 79-2 1 5LJ, rivulet exposure, no anneJid Jaws.

MALMS 1 . Hemse Beds, Hemse Mari, uppermost part and Eke Beds, lowest part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, b'"; Larsson 1 979; Cherns 1 983'". Samplesljawed annelids: Hemse Beds: 75-85CB, the uppermost Hemse bed, paulinitid jaws, annelid jaws; Eke Bads: 75-86CB, 0.20 m above the reference level, no annelid jaws.

MARTILLE 7. Slite Beds, unit d . Age: Early Wenlock. Reference: Jeppsson 1 983*. Samplesljawedannefids: 73- 1 8LJ, 0.05 m below soil suriace (not 0 .5 m , personal communication, Lennart Jeppsson; compare Jeppsson 1 983), fragments.

M I LLKLINT 3 . Hemse Beds, unit e (Mi l lkl int limestone). Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO; Jeppsson 1 974, 1 983; Laufeld & Jeppsson 1 976. Samp/es/jawed annelids: 69-30LJ, base of the section, sl ightly more than 1 m below the reference level , n o annelid jaws; 69-31 LJ , 0.50--0.60 m above base of section, no annelid jaws; 69-32LJ, 1 . 0--1 . 1 m above base of section, no annelid jaws; 69·35LJ, 2.50-2.65 m above base of section, no annelid jaws; 69-36LJ, annelid jaws; 69-37LJ, 3.5--3.6 m above base of section, no annelid jaws.

M OJ N E R 4. 639840 1 67882 (CJ 6904 9480), ca 2720 m ESE of Boge church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 69 Sl ite.

Small , shore-l ine exposure, about 400 m N E of the fishing huts (not marked on the topographical map sheet). There is a wide area with beach ridges outside of the fores1. Wheel tracks from this area join the small road that is marked on the topographical map. The exposure is beyond the beach ridges, nearly straight down to the shore from this intersection, at the southern end of the about 200 m long, reed-free shore. Slite Beds, Slite MarI. Samplesljawed anne/ids: 84-94LJ, shore exposure, annelid jaws.

MOLLBOS 1 . Halla Beds, unit b. Age: Late Wenlock. References: Laufeld 1 974a, bO ; Laufeld & Jeppsson 1 976; Claesson 1 979; Larsson 1 979; Laufeld & Martinsson 1 98 1 ; Uljedahl 1 98 1 , 1 983, 1 984, 1 985, 1 986; Stridsberg 1 98 1 a, b, 1 985; Jeppsson 1 983; Ramskåld 1 985b; Frykman 1 989. Samp/es/jawed annelids: 67-25LJ, diteh section, Kettnerites (K) bankvaetensis, K. (K) po/anensis, annelid jaws; 67-26LJ, ditch section, Kettnerites (K.) martinssonii, K (K) p% nensis, annelid jaws; 75- 1 6CB, about 20 m NW of the reference point and 0.40-0.45 m below the top of the section, annelid jaws; 75- 1 7CB, immediately above 75- 1 6CB, 0.30--0.40 m below the top of the section, Kettnerites (K.) bankvaetensis, K (K) martinssonii, annelid jaws; 77-28LJ, top of the section(?), Kettnerites (K.) bankvaetensis, K. (A) sisyphicf. var. val le, annelid jaws.

MOLLBOS 2 . Halla Beds, unit b. Age: Late Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979. Samples/jawedannefids: 79-81 LJ, Kettnerites (K.l p% nensis, annelid jaws.

MOLNER 1 . Mulde Beds, upper part. Age: Late Wenlock. Referenees: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 983. Samplesljawed annelids: 7 1 -6LJ, tap of the section, paulinitid fragments, annelid jaws; 7 1 -7LJ, 1 . 70 m below the top of the section and 0.70 m above the base of the section, Kettnerites (K.) martinssonii, K. (A.) sisyphi sisyphi, annelid jaws.

MULDE 2. Mulde Beds, uppermost part. Age: Late Wenlock.. References: Laufeld 1 974a, bO. Samples/jawed annelids: 75-26CB, middle of the ditch section, Kettnerites (K.) martinssoniivar. mulde, K. (K.) polonensis, K. (A) sisyphi sisyphi, annelid jaws.

MULDE TEGELBRUK 1 . Mulde Beds. Age: Late Wenlock. Referenees: Laufeld 1 974a, bO; Larsson 1 979. Samples/jawed annelids: 67-28LJ, loose slabs, Kettnerites (K) martinssonii var. mulde, K. (A) sisyphi sisyphi, annel id jaws; 82-7CB, loose slabs, Kettnerites (K.) martinssonii var. mulde, K. (K) polonensis, K (A) sisyphi sisyphi, K. (A) sisyphicf. val le , annelid jaws . .


M U N K EBOS 1 . Slite Bads, Slite Mari, Pentamerus gothlandicus Bads or slightly younger. Age: Wenlock. References: Laufeld 1 974a, b*, Samples/jawed annelids: 7 1 -84U, riv ulet section 25 m W of the reference pOint, Gotfandites slitensis, Hindenites gladiatus, Keltnerites (A.) sisyphi sisyphi, annelid jaws.

MYRSNE 1 . Slite Bads, Slite MarI. Age: Wenlock. Reference: Larsson 1 979*, Sampfes/jawed annelids: 82-43LJ, low ditch section, Kettnerites (K.) huberti, K. (K.) jacobi, K. (AJ sisyphi sisyphi, annelid jaws.

NABBAN 2. Eke Bads, lower part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, b*. Samples/jawed annelids: 72- 1 6LJ, shore exposure, same level as sample 72-60SL (Laufeld 1 974a), Kettnerites (K.) cf. bankvaetensis, K (K) martinssonii, annelid jaws.

NÅR 2 . Hemse Beds, Hemse MarI , uppermost part. Age: Ludlow, Late(?) Leintwardinian. References: Fredholm 1 988'. Samplesljawedannelids: 84-306DF, 0.82-0.78 m below the reference level, Lanceolatites gracifis, Kettnerites (K) martinssonii, K (K.) huberti, K (K) polonensis, annelid jaws; 82-326DF, 0.05--0 . 1 0 m above the reference leve l , Lanceolatites gracifis, Kettnerites (K.) martinssonii, K (K) huberti, annelid jaws.

NÅRS FYR 1 . Hamra Beds, lower part. Age: Late Ludlow, WhitcliHian. References: Laufeld t 974a, b-. Samplesljawedannelids:83-9GB, east of Nars Fyr 1 , extending the locality eastward, shore exposure in the southernmost part of Narsholmen immediately east of the small sea stack area, about 1 m above water level, close to a filIed fissure , no annelid jaws; 83-8CB, fissure f i l l ing, fragments.

NARSHAM N 2 . 634790 1 67246 leJ 5883 4503), ca 9480 m ESE of Burs church . Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 56 Ronehamn.

An exposure in the water, about 1 m from the beach in the shallow harbour between the two jetties, closer to the northern than the southern jetty. Burgsvik Beds, lower part(?). References: Hede 1 925a, the area i n general. Samples/jawed annelids: 83- 1 2LJ, shallow exposure, Kettnerites (K) martinssonii, K. (A) microdentatus, K. (K) polonensis, Hindenites naerensis, annelid jaws.

NARSHAMN 3 . 634770 1 6721 8 leJ 5852 4482), ca 9490 m ESE of Burs church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 56 Ronehamn.

Shore exposure with low sections with fossil wave marks, where the road is at its closest to the sea, about 200 m south of the southernmost jetty at Narshamn. Burgsviks Beds, lower part(?). References: Hede 1 925a, p . 3 1 , the area i n general . Samplesljawed annelids: 83- 1 3LJ, above an uneven erosion surface, Kettnerites(K.) cf. martinssonii, K. (A.) microdentatus, annelid jaws.

NISSE 1 . Hemse Beds, Hemse Mari NW part. Age: Ludlow. References: Laufeld 1 974a, bO; Larsson 1 979. Samples/jawed annelids: 75-98CB, in a water hole close to the ditch, nodular lutitic calcarenite in soft mari, Kettnerites (K) huberti, annelid jaws.

NORRVANGE 1 Slite Beds, unit a. Age: Early Wenlock. References: Laufeld 1 974a, bO. Samples/jawed annelids: 82-46CB, 0.65 m below the top of the inland cl iff section, annelid jaws; 82-47CB, about 0 .2 m below 82-46CB, no annelid jaws.

NORS t . Hogklint Beds, unit b upper part. Age: Early Wenlock. References: Laufeld 1 974a, b-. Samples/jawedannelids: 82-50CB, uppermost exposed bed at the strand, no annelid jaws.

NORS STENBROTT 1 . 642580 1 6851 4 leK 7740 21 66), ca 6230 m N N E of Fleringe church. Topographical map sheet 7 J Fårosund SO & NO. Geological map sheet Aa 1 71 Kappelshamn.

Quarry, situated al the end of the small road marked with red on the map. The quarry, which is not marked on the 2nd edition of the topographical map, is marked on the 3rd edition. The section is immediately SE of the entranee of the quarry. Reference point: The easternmost point of the quarry close to and south of the entrance. Reference level: The contact batween the lowe� brownish-grey, thin-bedded calcarenite and the light-coloured, more thickly bedded calcarenite, about 1 .85 m above the base of the quarry. Hogklint Beds, unit b(?) . Age:Wenlock, Sheinwoodian. Samples/jawedannelids: 8249GB, about 1 .6 m above the base of

. the section and 0.2-0.25 m below the very distinet contact between the light and the dark calcarenite, Kettnerites (K.) abraham isaac, K (K.) aH. bankvaetensis, annelid jaws.

NYAN 1 . Eke Beds, lower part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979. Sampleljawed annelids: 72-9LJ, excavated material, no annelid jaws; 72- 1 0LJ, excavated material, Kettnerites (K) cf. bankvaetensis, annelid jaws; 75-63CB, excavated material, pauli nitid jaw.

NY AN 2 . Hemse Beds, uppermost part and Eke Beds, lowest part. Age: Ludlow, Late Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979; Cherns 1 983°; Fredholm 1 988.


Samples/jawed annelids: Hemse Beds: 72- 1 4LJ , 0.78-0 . 8 1 m below the reference level, fragments; 72- 1 3LJ, 0.45--0.51 m below the reference level, Kettnerites (K) huberti, annelid jaws; 72- 1 2LJ, 0-0 . 1 0 m below the reference level, Kettnerites (K) huberti, K (K) cf. polonensis, annelid jaws; Eke Beds: 72- 1 1 LJ, 0-0.21 m above the reference level, paul in itid jaws, annelid jaws; 75-62GB, uppermost bed in the section, no annelid jaws.

NYGARDS 1 . Slite Beds, Sl ite Mari, Pentamerus gothlandicus Beds. Age:Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979; Ramskold 1 983. Samples/jawed annelids: 75- 1 4CB, about 0 .75 m below ground level, Kettnerites (Kl martinssonii, K (A) sisyphi sisyphi, Gotlandites slitensis, annelid jaws.

NYGARDS 2 . Halla Beds , unit b . Age: Late(?) Wenlock. References: Laufeld 1 974a, bO. Sampfesljawed annelids: 75- 1 5CB, loose slabs, Kettnerites (K) bankvaetensis, K. (A.) sisyphi sisyphi, annelid jaws; 8 1 -75GB, shallow ditch, Kettnerites (K) bankvaetensis, K (K.) martinssonii, K (K) polonensis, Hindenites gladiatus, annelid jaws.

NYGARDSBAcKPROFILEN 1 . Lower and Upper Visby Beds. Age: Early Wenlock. Reference: Jeppsson 1 983°. Samplesljawed annelids: Lower Visby Beds, unit e: 8 1 -2LJ, 0.0 m a . s . l . , Kettnerites (K) martinssonii, K (K.) abraham abraham, K. (K) versabilis, annelid jaws; 79-42LJ, about 2.25 m a .s . ! . , K. (K) versabilis, K (K) versabilis C form, Lanceolatites gracilis, Kettnerites (K) abraham abraham, annelid jaws; Upper Visby Beds: 79-43W , about 4.6 m a .s . l . , Kettnerites (K.) abraham abraham, annelid jaws.

NYHAMN 1 . Lower Visby Beds, unit b. Age: Late Llandovery to Early Wenlock. Reference: Laufeld 1974a, bO, Larsson 1 979. Samplesljawedannelids: 79-21 7LJ, about 2.5 m a.s. l . , Kettnerites (K) versabilis, K. (K.) abraham abraham, annelid jaws; 8 1 -28LJ, about 1 .5 m a .s . l . , jaw fragments.

NYHAMN 2 . Lower Visby Beds unit b. Age: Late Llandovery to Early Wenlock. References: Laufeld 1 974b", Larsson 1979. Samplesljawedannelids: 75-2CB, shore exposure, Kettnerites (K) abraham abraham, annelid jaws.

NYHAMN 4. 640700 1 65521 leK 461 5 0523), ca 2720 m WSW of Lummelunda church. Topographical map sheet 7 J Fårosund SV & NV. Geological map sheet Aa 1 83 Visby & Lummelunda.

Shore exposure 40-50 m N of the bay and 5 m N of the smal l clearanee in the beach cobble made to permit small boats to be dragged up on land. The locality is between the western most, medium-sized lichen-covered boulders on the shore. Lower Visby Beds, unit b. Samplesljawed annelids: 79- 1 4LJ, surface exposure, Kettnerites (K) versabilis, K. (K.) martinssonii, K (K) abraham abraham, Kettnerites sp. , annelid jaws.

NYHAMN 5 . 640620 1 655 1 0 leK 4592 0445), ca 3240 m SW of Lummelunda church. Topographical map sheet 7 J Fårosund SV & NV. Geological map sheet Aa 1 83 Visby & Lummelunda.

Small shore exposure between erratic boulders. The locality is marked on the geological map about 1 000 m S of Nyham n. Lower Visby Beds, unit b . Samplesljawed annefids: 84- 1 1 LJ, shore exposure, Kettnerites (A.) microdentatus, Kettnerites sp. , annelid jaws.

NYMANETORP 1 . Hogklint Beds, unit b . Age: Early Wenlock. References: Laufeld 1 974a, bO; Laufeld & Martinsson 1 98 1 . Samplesljawed annefids: 81 -64 W , 2.0 m below the reference leve l , no annelid jaws.

OIVIDE 1 . Sl ite Beds, unit f (Rhipidium tenuistriatum Beds). Age: Early Wenlock. References: Laufeld 1 974a, b-; Larsson 1 979; Jeppsson 1 982, 1 983. Samplesljawed annelids: 79-44 W , 0.5 m below the tap of the section, Kettnerites fragment, annelid jaws.

OLLAJVS 1 . Hamra Beds, unit c . Age: Late Ludlow. References: Laufeld 1 974a, bO. Sampfesljawed annelids: 76- 1 7GB, ditch section, about 0.5 m above the bottom of the ditch, Kettnerites (Kl cf. bankvaetensis, annelid jaws.

OLSVENNE 3. Eke Beds, lowest part. Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, bO. Samplesljawed annelids: MS90 1 AM, paul in itid sp. , annelid jaws; 74-32KL, Kettnerites sp., annelid jaws.

ONDARVE 2 . Hemse Beds, Hamse Mari, uppermost part, Eke, base. Age: Ludlow, Late Leintwardinian . Reference: Fredholm 1 988. Samplesljawed annelids: Hemse Beds: 83-5LJ, ditch exposure, Kettnerites (K.) martinssonii, K. (K.l huberti, annelid jaws.

OSTERGARNSHOLM 1 . Hemse Beds, unit e(?). Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, b-; Larsson 1 979; Jeppsson 1 983. Samples/jawed annelids: 78-1 9CB, about 0.50 m above the base of the section which is covered by loose gravel, no annelid jaws.

OSTERGARNSHOLM 2 . Hemse Beds, unit e(?). Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 982; Fred holm 1 988.

Sample/jawed annefids: 78- 1 8GB, 0 .6 m above the flat surface, just above sea level, no annelid jaws.

OSTERGARNSHOLM 3 . Hemse Beds, unit et?). Age: Ludlow, Leintwardinian. Reference: Jeppsson 1 983; Fredholm 1 988°. Sampleljawedannelids: 78-20CB, middle part of the shallow shore exposure, no annelid jaws.

OTES 1 . Sundre Beds, middle part. Age: Latest Ludlow. References: Laufeld 1 974a, bO. Samples/jawed annelids: 75- 1 6 LJ, about 200 m S of the monument, no annelid jaws.

PETSARVE 2 . Eke Beds, middle-upper part. Age: Ludlow. References: Laufeld 1 974a, b·. Sampfes/jawedannelids: 77-34CB, excavated material from ditch, Lanceolatites gracilis, Kettnerites (K.) martinssonii, K. (K) huberti, annelid jaws.

PETSARVE 1 5 . Eke Beds, middle-upper part. Age: Late Ludlow(?). References: Laufeld 1 974a, bO; Larsson 1 979. Samples/jawedannelids: 77-33GB, bottom of the ditch, Kettnerites (K) martinssonii, K (Kl huberti, annelid jaws.

PRAsTBATELS 2 . Klinteberg Beds, lower part. Age: Latest Wenlock. References: Laufeld 1 974a, bO, Frykman 1 989. Samples/jawed annelids: 75-93CB, 1 .20 m below the tap of the section and about 0.3-0.5 m above the base of the section, n o annelid jaws; 75-94CB, 0.85 m above 75-93CB, no annelid jaws; 75-95CB, loose stab, no annelid jaws.

RAGAKRE 1 . Slite Beds, Slite Siltstone, Halla Beds and Klinteberg Beds, lower part. Age: Late Wenlock. References: Laufeld 1 974a, bO; Sivhed 1 976 (Dans 2); Bergman 1 979b, 1 980a; Frykman 1 989. Samples/jawed annelids: 76-3 1 GB, from the limestone with wave marks, rich i n bryozoans, at the base of the quarry, Kettnerites (K) martinssonii, annelid jaws; 76-28C B, softcalcilutite above the wave marks in the floor of the quarry, Kettnerites (K) cf. martinssonii, K (K.) cf. po/onensis, annelid jaws.

RANGSARVE 1 . Hemse Beds, upper part. Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, bO; Laufeld & Jeppsson 1 976; Claesson 1 979; Larsson 1 979; Laufeld & Martinsson 1 98 1 ; Ramskold 1 983; Jeppsson 1 983; Fredholm 1 988. Samplesljawed annelids: 71-1 06LJ, base of the section, a more than 0.35 m thick bed, fragments; 7 1 - 1 07LJ, 0.35--0.40 m above the basal bed, fragments; 7 1 - 1 08LJ, 1 .00-1 .08 m above the base of the section, Kettnerites (K) cf. martinssonii, K. (A) sisyphi, annelid jaws; 7 1 - 1 09LJ, 2.28 m above the base of the section, annelid jaws; 7 1 - 1 1 OLJ, 3. 1 8 m above the base of the section, fragmented paulinitid jaws, annelid jaws; 75-59CB, 0.5 m above road surface, Kettnerites (A) d. sisyphi, annelid jaws; 75-58CB, about 1 .75 m above road surface, no annelid jaws; 75-57GB, about 3.0 m above road surface, no annelid jaws; 75-?6CB, top of the section, 4.5 m above road surface, no annelid jaws.

ROBBJÅNS KVARN 2. Slite Beds, Slite Siltstone, and Halla Beds. Age: Wenlock. References: Laufeld 1 974a, bO; Sivhed 1 976; Bergman 1 979b, 1 980a.

•

Samples/jawedannelids: Halla Beds: 7 1 - 1 39LJ, at the east-southeasternmost extension of the locality, annelid jaws.

ROBBJANS KVARN 3. 636361 1 64527 leJ 3294 6274), ca 820 m SW of Kl inte church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 60 Kl intehamn.

Ditch exposure ca 100 m E of the house at Robbjans Kvam 2. Mulde Beds, lower part. Age: Late Wenlock. Samples/jawed annelids: 7 1 - 1 38LJ, ditch exposure, Kettnerites (A) sisyphi sisyphi, annelid jaws.

RONEHAMN 1 . Eke Beds and Burgsvik Beds. Age: Late Ludlow. References: Laufeld 1 974a, bO. Samples/jawedannelids: Eke Beds: 75-84CB, loose slabs of excavated material about 1 5 m N of the northernmost oil storage tank, annelid jaws.

RONEHAMN 2 . Eke Beds and Burgsvik Beds. Age: Late Ludlow. Reference: Larsson 1 979. Samplesljawed annelids: Eke Beds: 75-24LJ, exposure on the shore, Kettnerites (K.) polonensis, annelid jaws; 75-25LJ, loose material, no annelid jaws.

RONNINGS 1 . Eke Beds, upper part. Age: Ludlow, Whitcliffian. References: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 983. Samplesljawedannefids: 7 1 - 1 90LJ, uppermost exposed bed, Kettnerites (K.) martinssonii, K (K.) hubert;, annelid jaws; 75-26LJ, 0.2 m below 7 1 - 1 90 , Kettnerltes (K) martinssonii, annelid jaws.

RONNKLINT 1 . Lower Visby Beds, Upper Visby Beds, and Hogkl int Beds. Age: Late Llandovery to Early Wenlock. Referenees: Hede 1 940, p. 13 l ines 1 6- 1 7 ; Brood 1 982, 1 985; Jeppsson 1 983·; Ramskold 1 984, 1 985b. Samples/jawedannelids: Lower Visby Beds, unit b : 8 1 - 1 1 LJ, about 7.6 m below the reference level at the sea level, Lanceolatites gracifis, Kettnerites (K.) abraham abraham, K (A) siae/soeensis, annelid jaws; 79-2 1 1 LJ, about 6 . 1 0-6.30 m below the reference level, Kettnerites (K.) martinssonii, K. (K.) abraham abraham, annelid jaws; 79-2 1 2LJ, 4.05-4.00 m below the reference level, Kettnerites (K) martinssonii, K. (K) abraham abraham, annelid jaws; 79-21 3LJ, 2 .05-1 .95 m below the reference level, no annelid jaws; 79-2 1 4LJ, 0.05--0.08 m below the reference level, Kettnerites


(A.) sisyphi sisyphi, K. (A.) siaelsoeensis, K. (K.) abraham abraham, annelid jaws; 8 1 -24LJ , 0.25-0.30 m above the reference level, Ketlnerites (K.) abraham abraham, annelid jaws.

RUDVIER 1 . Hamse Bads, unit et?) and d. Age: Ludlow. References: Mori 1 970' Ioc. 1 1 6 ; Ramskold 1 986*; Fredholm 1 988. Samples!}awedannelids: Hamse Bads, unit et?): 84-3 1 0DF, from a lens of dense marly l i mestone within the shale, Kettnerites (K.) polonensis, annelid jaws.

SAXRIV l. Hogklint Bads unit b. Age: Early Wenlock. Reference: Hede 1 933':23 l ines 22 to 30, page 24 l ines 1 -1 5 ; Larssen 1 979", Reference point:The boulders leaning against the section sean on the photograph (Hede 1 933, p. 25), the boundary of Hede's laeal units a-b. Referenee /eve/:The deepest eroded bed about 0.5 m above water level. Samp/esljawed annelids: 84-77LJ, 0.45--0.40 m below the reference level, Kettnerites (K.) martinssonii, K. (K.) cf. abraham, K. (A) sisyphi sisyphi, Hindenites gladiatus, anne lid jaws; 84-79LJ, 3.8 m above the reference level, Hede's local unit c, fragments.

SI BBJANS 2. Hamra Beds, unit b. Age: Latest Ludlow. Referenees: Laufeld 1 974a, b-. Samp/esljawedannelids: 82-32LJ, bottom of the ditch , Kettnerites (K.) p% nensis, annelid jaws.

SIGVALDE 2. Hemse Beds, lower-middle (probably unit c, perhaps also unit dl. Age: Early(?) Ludlow. Referenee:Jeppsson 1 983·; Fredholm 1 988. Reference /evel: The upper surface of the /I{onja prisca bed (Hede 1 925a : 1 7). Reference point: The fissure about 30 mm from the left side of the photograph i n Munthe ( 1 9 1 0) and Hede ( 1 925a, p. 1 7) . Samp/es/jawed anne/ids: 7 1 - 1 1 6LJ, 0.95--0.90 m below t h e reference level , Kettnerites (K.) martinssonii, K. (K.) cf. pa/onensis, annelid jaws; 7 1 - 1 1 5LJ, the /lionia prisca bed, Kettnerites (K.) martinssonii, K. (K.) pa/anensis, K (A) sisyphi sisyphi, annelid jaws; 7 1 - 1 1 7LJ, 0.70-0.80 m above the reference level and adjacent to the reference point, fragments.

SION 1 . Slite Beds, unit f ( Rhipidium tenuistriatum Beds). Age: Early Wenlock. References: Laufeld 1 974a, bO; Jeppsson 1 983. Samp/esljawed annelids: 72-3LJ, shaliow exposure, same sample level as 72-46SL (Laufeld 1 974a), no annelid jaws.

SJALS6 1 . Lower Visby Beds , unit b. Age:Late LIandovery. Reference: Larsson 1 979*. Samp/es/jawed anne/ids: 79- 1 2LJ, surface exposure, Kettnerites (K.) abraham abraham, K. (A.) siae/soeensis, annelid jaws.

SJALSO 3 . 639920 1 65 1 90 (CJ 4227 9770), 4200 m W of Våskinde church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 83 Visby & Lummelu nda.

Bottom of the shallow harbour beside the jetty marked with the year 1 970. Lower Visby Beds, unit b (?). Age: Late LIandovery. Samples/jawed annelfds: 8 1 -55LJ, topmost bed at the outer end of the jetty, annelid jaws.

SJAUSTR EHAMMAR l. 636453 1 68056 (CJ 681 5 6097). ca. 3770 m S of Gammelgam church. Topographical map sheet 6 J Roma SO. Geological map sheet Aa 1 70 Katthammarsvik.

Gliff section on the southern part of the peninsula Sjaustrehammar, SW of the triangulation point. The locality is shown on the photograph i n Hede 1 929, p. 45. Hemse Beds. unit d . Age: Ludlow. Reference: Hede 1 929, pp. 44-45. Samples/jawed annelids: 82- 1 8LJ, 0.5 m below the top of the distinet, thinly bedded caJcilutite, no annelid jaws; 82- 1 9LJ, 0.5 m above the thinly bedded calcilutitic bed, Kettnerites (K.) polanensis var. sjaustre, Kettnerites (K.) sp. , anneli d j aws ; 82-20LJ, the Mega/amus l imestone below the thinly bedded calcilutitic bed, no annelid jaws.

SKALASANDSVIK 2 . 643068 1 70803 (DK 0045 2479), ca 1 1 680 m ENE of Fåre church . Topographical map sheet 7 K Ul lahau NV. Geological map sheet Aa 1 80 Fåre.

Shore section about 1 340 m E of the triangulation point 22,3 at Lansa nåsskifte, the area with ripple marks and west of them. Slite Beds, Slite MarI. Age: Wenlock. Reference leve/: The upper surface of the wave marks. References: Hede 1 936, p. 33; Bergman 1 979b, p. 221 , Ioc. 6 and 7. Samplesljawed annelids: 80-5GB, 0.5 m above the wave mark level and 40 m W of the westernmost wave marks, fragments.

SKENALOEN L 640004 1 68599 (CJ 7628 9595). ca 9950 m SSE of Hellvi church:Topographical map sheet 7 J Fårosund SO & NO. Geological map sheet Aa 1 69 Slite.

Shore exposure on the southern point of the island Skenalden about 1 m above sea level. Slite Beds, unit g . Age:Wenlock. Samplesljawed annelids: 75-6GB, argillaceous calcarenite, rich in stromatoporoids, annelid jaws.

SKENALOEN 2 . 64001 6 1 68603 (CJ 7633 9607), ca 9850 m SSE of Hellvi church. Topographical map sheet 7 J Fårosund SO & NO. Geological map sheet Aa 1 69 Slite.

Cliff section on the eastern shore of the island of Skenalden. Reference point:The ca 1 m wide, dike-like structure composed of argiliaceous material darker than the surrounding limestone. Slite Beds, unit g . Age: Wenlock.

Samples/jawedannelids: 75-7CB, from the lower part of the structure, no annelid jaws; 75-8GB, about 1 m above previous sample and from the upper part of the section, paul in itid jaws, annelid jaws.

SLA THALLAR 3. Hogklint Beds, middle part. Age: Early Wenlock. Reference: Larsson 1 979. Samp/esljawedannelids: 75-42KL. annelid jaws.

SlITEBROTTET 1 and 2. Slite Beds, Sl ite Mari and Slite g . Age: Wenlock. References: Laufeld 1 974a, b*; Walmsley & Boucot 1 975, p. 65; Eisenack 1 975, Fig. 1 8 ; Larsson 1 979; Jeppsson 1 983'; Odin et al . 1 984, 1 986. Samplesljawedannelids: SLITEB ROTTET 1 (ali levels are given in relation to the top of the section, except those referring to marker leveis) : 73-32LJ, 0 . 1 5 m, Kettnerites (K.) martinssonii, K. (A) sisyphi sisyphi, Gotlandites slitensis, Hindenites sp. , annelid jaws; 73-33LJ, 2.0 m, Gotlandites sp., annelid jaws; 73-34LJ, 4.40 m below the top of the section and 1 . 20 m above a 0.37 m thick, deeply eroded marly, level which can be followed around the quarry, Kettnerites (K.) martinssonii, K. (A) sisyphi sisyphi, K. (K.) cf. jacobi, Gotlandites slitensis, Hindenites sp., annelid jaws; 73-36LJ, 8.55 m below the top of the section, immediately below a 0.20 m , deeply eroded level and 0.50 m above another deeply eroded level (0.23 m thick), no annelid jaws; 73-37LJ, 1 0.45 m, Kettnerites (K.) bankvaetensis, annelid jaws; 73-38LJ, 1 1 .45 m below the top of the section and 0 045 m below a dril led drainage hole in the section, Kettnerites (K.) martinssonii, K (A) sisyphi sisyphi, annelid jaws; 73-39LJ, 1 3 .60 m fragments; 73-40LJ, 1 5 .70 m , Kettnerites (K.) martinssonii, K. (A) sisyphisisyphi, Gotlandites slitensis, annelid jaws; 73-4 1 LJ , 1 8.60 m, Kettnerites (K.) bankvaetensis, K. (K) martinssonii, K. (A) sisyphi sisyphi, annelid jaws; 73-42LJ, 20.90 m, Kettnerites (K.) martinssonii, K. (A) sisyphi sisyphi, Hindenites sp. , anne l id jaws; 73-44LJ, 25.60 m , no annelid jaws. SLITEBROTTET 2 (Ieveis are related 10 the top of the section measured from Slitebrottet 1 and thus forming a continuous section): 83-31 LJ, 35.90 m below the top of the section and 0.00-0 . 1 0 m above the lower reference level, Kettnerites (K.) martinssonii. K. (K.) jacobi, K (K.) pofonensis, K (A) cf. microdentatus, Hindenites el. gladiatus, annelid jaws; 83-32LJ, 36. 1 0 m below the top of the section and 0.00-0.20 m below the lower reference level, annelid jaws; 83-33LJ, 37.70 m below the top of the section and 1 .67-1 .70 m below the lower reference level, Kettnerites {K.) martinssonii, K. (K.) jacobi, K. (A) cf. sisyphi, annelid jaws; 83-34, 40.60 m , Kettnerites (K.) martinssonii, K. (K.) jacobi, K. (A) cf. microdentatus, anneIld jaws; 73-48LJ, 40.80 m . Kettnerites (K) martinssonii, K (K.) jacobi, K. (A) sisyphi sisyphi, K. (A) mieradentatus, annelid jaws; 83-35LJ, 41 .70 m, Kettnerites (K.) martinssonii, K. (K.) jacobi, K. (A) sisyphi sisyphi annelid jaws; 73-49LJ, 42.90 m , pau l in itid fragments, annelid jaws; 83-36LJ, 43.95 m, Kettnerites (K.) martinssonii, K. (K.) jacobi, K. (A) sisyphi sisyphi, Hindenites gladiatus, annelid jaws; 84-1 LJ, 49.60 m, Kettnerites (K.) jacobi, K. (A) sisyphi sisyphi, annelid jaws.

SMISS 1 . Klinteberg Beds, Klinteberg Mari, top. Age: Ludlow. References: Laufeld 1 974a, b�; Larsson 1 979; Frykman 1 989. Samples/jawedannelids:75-76GB, loose slabs from the temporary excavation, Kettnerites (A) microdentatus, annelid jaws.

SNACK l . 639670 1 65075 (CJ4096 9529). ca 5900 m WSW of Vaskinde church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 83 Visby & Lummelunda.

Temporary excavation, about 1 00 m NW of the northwesternmost corner of the main bu i lding of Snack, not marked on the topographical map. Lower Visby Beds. Age: Late LIandovery. Samp/es/jawed annelids: 83-1 GB, about 1 .5 m below the road surlace, Kettnerites (A) cf. microdentatus, annelid jaws.

SNAGKGARDSBADEN 1 . Upper Visby Beds and Hegklint Beds, unit a. Age: Early Wenlock. References: Laufeld 1 974a, b*, Larsson 1 979. Samples/jawedannelids: Upper Visby Beds: 79-22LJ, 0.5 m above road surface, lowest part of the section, no annelid jaws; 79-25LJ, 6.3 m above sample 79-22, probably about 1 .0 m below the reference level, Lanceolatites gracilis, Kettnerites (K.l martinssonii, K. (K.l abraham abraham, annelid jaws; 79-26LJ, about 2 m below the reference leve l , Kettnerites (K.l abraham abraham, annelid jaws; Hegklint Beds, unit a: 79-27LJ, lowest Hegklint Beds, unit a. the reference level, Kettnerites (K.) martinssonii, annelid jaws; 79-28LJ, about 2 m above the reference level , Kettnerites (K.l martinssonii, K. (A) sisyphi, anneI ld jaws.

SNAUVALDS 1 . Hemse Beds, Hemse Mari NW part. Age:Ludlow, Bringewoodian or more probably Early Leintwardinian. Referenees: Laufeld 1 974a, b*; Larsson 1 979; Jeppsson 1 983; Fred holm 1 988. Samplesljawed annelids: 7 1 - 1 4LJ, ditch section, Kettnerites (K.) huberti, K. (A) sisyphi sisyphi, annelid jaws.

SNODER 2. Hemse Beds, Hemse Mari NW part. Age: Ludlow. References: Laufeld 1 974a, b"; Larsson 1 979; Ramskold 1 983, 1 984, 1 985b. Samplesljawed annelids: 82- 1 4CB, loose slab from the drainage ditch, Lanceo/atites gracifis, Kettnerites (K) martinssonii var. mulde, K. (A) sisyphi sisyphi, K. (A) fjae/ensis, annelid jaws.

SNODER 3. Hemse Beds, Hemse Mari NW part. Age: Ludlow, Bringewoodian or Early Leintwardinian. Reference:Jeppsson 1 983*; Fredholm 1 988. Samp/es/jawed annelids: 7 1 - 1 OLJ, bottom of the small ditch, Kettnerites (K.) martinssonii var. mulde, K. (A) cf. sisyphi, anne l id Jaws.

SOORA TORG , . 639279 1 64840 (CJ 3828 9 1 60), ca 800 SW of Visby cathedral. T opographical map sheet 6 1 Visby NO. Geological map sheet Aa 1 83 Visby & Lummelunda.


Temporary excavation along t h e western side o f t h e marke1 place (Sedra Torg) , about 28-29 m a.s. 1 . Hogklint Beds, unit c . Age: Early Wenlock. Samplesljawedanne/ids: 75-6LJ, about 0.75 m below the top of the section, no annelid jaws; 75-7LJ, about 0.7 m above 75-6, no annel id jaws.

SOJVIDE 1 . Slite Beds, unit f (Rhipidium tenuistriatum Beds). Age: Early Wenlock. References: Larsson 1 979*; Jeppsson 1 983. Samples/jawed annelids: 8 1 -58LJ , very small ditch exposure, Kettnerites (A) sisyphi sisyphi, pauli nitid sp. , annelid jaws.

SPROGE 3. 635024 1 64538 (CJ 3203 4960) , ca 280 m N E of Sproge church. Topographical map sheet 61 Visby SO. Geological map sheet Aa 1 64 Hemse.

Ditch exposure along and S E of the road between Levide and Sproge, 25 m N E of the minor road towards the SE and about 225 m E of the main rcad ( 1 40). Hemse Beds, Hemse Mari NW part. Age: Ludlow. Samples/jawed annelids: 79-9LJ , bottom of the ditch, no annelid jaws.

SPROGE 4. 6350 1 6 1 64434 (CJ 3097 4940), ca 840 m W of Sproge church. Topographical map sheet 61 Visby SO. Geological map sheet Aa 1 64 Hemse.

Ditch exposure along and south of the road, about 1 00 m W of the farm house at Alvegårde, just E of the entrance to the tield. Hemse Beds, Hemse Mari NW part. Age: Ludlow. Samp/es/jawedannelids: 84-80LJ, bottom of the ditch, Kettnerites (K) martinssonii var. mu lde, K. (A) sisyphi sisyphi. K. (A) microdentatus, K. (A) fjae/ensis, annelid jaws.

SPROGE 5. 634984 1 64382 (CJ 3045 49 1 0) , ca 1 380 m WSW of Sproge church. Topographical map sheet 5 I Hoburgen N O & 5 J Hemse NV, Geological map sheet Aa 1 64 Hemse.

Road-side ditch exposure E of the rcad WSW of Alvegårde, about 200 m N of the small road leading to the fann SSW of Alvegårde. The locality is where a small ditch runs from a bushy area east of the road and meets the main ditch beside the road. Hemse Beds: Hemse Mari NW part. Age: Ludlow. Samples/jawed annelids: 84-81 LJ, 0.5 m above the bottom of the ditch, annelid jaws.

STAVE 1 . Sl ite Beds, Slite Mari, central part. Age: Early Wenlock. References: Laufeld 1 974a, b-; Larsson 1 979. Samples/jawed annelids: 75- 1 1 GB, 0.5 m below ground surface, Kettnerites (K) martinssonii, K. (K.) cf. jacobi, K. (A) microdentatus, annelid jaws.

STENSTU 2. 637539 1 6 7433 (CJ 6278 7224), ca 1 1 50 m SSE of Anga church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 70 Katthammarsvik.

Ditch exposure W of the old road at point 1 2 ,8; note that the rcad has now been rebuilt. Klinteberg Beds, unit f. Age: Early Ludlow. Samplesljawedannelids: 75-39LJ, ditch exposure, fragments.

STENSTUGÅRDS 1 . Klinteberg Beds, lower part. Age: Latest Wenlock. References: Laufeld 1 974a, b*. Samp/esljawed anne/ids: 78-7GB , 0 . 1 5 m below the top of the section in the SE corner of the quarry, no annelid jaws.

STORA BAN N E 2. Slite Beds, unit g , lower part. Age: Wenlock. Reference: Laufeld 1 974b·. Samp/es/jawed annelids: 69-1 LJ, 0040-0.50 m above the base of the section, no annelid jaws.

STORA BANNE 3. Slite Beds, unit g , lower part. Age: Wenlock. Reference: Laufeld 1 974b·. Samples/jawed annelids: 69-2LJ, 1 .40 m below the top of the section, no annelid jaws; 69-3LJ. 3.55 m below the top of the section, no annelid jaws.

STORA KRUSE l . 634487 1 653 1 1 (CJ 3932 4343), ca 1 380 m WSW of Alva church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 64 Hemse.

Ditch, 25 m E of the eas1ernmost house (not marked on the topographical map sheet) at Store Kruse, a ditch running N-S along the boundary between the two fields, north of and by the road. Hemse Beds, Hemse Mari S E part. Age: Ludlow. Samplesljawed annelids: 82-29LJ, loose slabs within the ditch , Lanceolatites graci/is, Langeites glaber, Kettnerites (A) sisyphi klasaardensis, K. (K) huberti, annelid jaws.

STORA MAFRIOS 2. 637287 1 64225 (CJ 3060 7221 ), ca 2500 m NW of Våstergarn church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 60 Kli ntehamn .

Ditch exposure, 1 5 t o 20 m NW o f the road from t h e small ditch that runs NW to Idån. Slite Beds, unit f, Rhipidium tenuistriatum zone or sl ightly younger. Samplesljawedannelids: 84-67LJ, excavated material, Kettnerites (K.) jacobi, K. (A) microdentatus, annelid jaws.

STORA MYRE 1 . Slite Beds, unit d . Age: Early Wenlock. References: Laufeld 1 974a, b-; Larsson 1 979; Jeppsson 1 983. Samp/es/jawed annelids: 7 1 -74LJ, surface exposure, Kettnerites (K.) martinssonii, K. (K.) bankvaetensis, annelid jaws; 75-29GB, surlace exposure, Kettnerites (K.) martinssonii, K. (A) sisyphi sisyphi, anne lid jaws.

STORA TUNE 1 . Klinteberg Beds, lower part. Age: Latest Wenlock. References: Laufeld 1 974a, b*; Frykman 1 989.


Samples/jawed annefids: 78-8CB, a surtace exposure SE o f the building and 3 m from the road, no annelid jaws.

STORA VIKARE 2. Halla Bads, unit b. Age: Late Wenlock. References: Laufeld 1 974a, bO, Samples/jawed annelids: 75- 1 8CB, shallow excavation, Kettnerifes (K.) polonensis, K. (A.) sisyphi sisyphi, annelid jaws.

STORUGNS , . Slite Bads, unit e (Kalbjerga Limestone). Age: Early Wenlock. Reference: Laufeld 1 974b*. Sampfesljawedannelids: 78-1 CB, about 1 . 5 m above the base of the section, no annelid jaws; 78-2CB, about 6 m above the base of the section, crinoid- and stromatoporoid-rich l imestone immediately abolla a homogeneous white l imestone, no annelid jaws; 69-1 6LJ, no annelid jaws.

STORUGNS l B. 641 572 1 679 1 5 (CK 7058 1 2 1 7) , ca 5 1 00 m N N E o f Lårbro church. Topographical map sheet 7 J Får6sund 80 & NO. Geological map sheet Aa 1 7 1 Kappelshamn.

Part of the large quarry Storugns, ca 320 m NNE of point 28,3 at Storugns and aboul 1 60 m E of Storugns 1 , where the road from Klinthagen quarry, S of Storugns, enters the Quarry. Reference level: A marker level high up on the walls can be followed around a large part of the Quarry. It consists of 0.52 m argillaceous limestone with severai pyrite horizons. The lowar boundary consists of a bed up to 1 0 mm thick, with pyrite, the base of which is the reference level. Thera are also a number of higher pyrite lave Is, up to 1 . 1 0 m above the reference leveL Slite Beds, u nits e and g. Age:Wenlock. Samplesljawed annelids: 84-29LJ, 0.01 -0.20 m above the reference level, Kettnerites (K.) bankvaetensis, annelid jaws; 84-28U, 0.32-0.52 m above the reference level, Kettnerites (K.) bankvaetensis, K (K) martinssonii, annelid jaws.

STRANDAKERSVIKEN 1 . Hegklint Beds, unit c and Slite Beds, unit c. Age: Early Wenlock. References: Laufeld 1 974a, bO. Samplesljawed annefids: Hegklint Beds, unit c: 77- 2 1 G B , about 0.4 m below the reference level, Kettnerites (K) martinssonii, K (K.) polanensis, Hindenites angustus, annelid jaws. Slite Beds: 77-22GB, 0 . 1 -0 . 2 m above the reference level, no anne lid jaws.

STRANDS 1 . Hamra Beds, unit b. Age: Latest Ludlow. References: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 983. Samples/jawed annelids: 75- 1 4U, bottom of the ditch, Lanceofatites sp. A, Kettnerites (K.) bankvaetensis, K. (K.) huberti, K. (K.) cf. polonensis, annelid jaws; 76-1 9GB, bottom of the ditch , Lanceofatites sp. A, Kettnerites (K) bankvaetensis, K. (K.) huberti, annelid jaws.

STUTSVIKEN 1 . Hogklint Beds unit c. Age: Early Wenlock. References: LaufeJd 1 974a, bO. Samplesljawed anne/ids: 77-20GB, Kettnerites (K) martinssonii, K (K) abraham isaac, annelid jaws.

SUDERBYS 3. 638740 1 67425 (CJ 6356 8423), ca 1 050 m WNW of Gothem church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 69 Sl ite.

Ditch exposure, directly north of Suderbys in the ditch along the road and about 1 m W of the N-8 ditch. Klinteberg Beds, unit b or c . Age: Late Wenlock. Reference: Hede 1 928, p. 58, l ines 28-3 1 . Sampfes/jawedannelids: 75-35LJ, ditch exposure, Kettnerites (K.) bankvaetensis, annelid jaw fragments.

SUDEAS 1 . Su ndre Beds, middle part. Age: Latest Ludlow. References: Laufeld 1 974a, bO. Samples/jawed annelids: 75- 1 8U , bottom of the ditch, annelid jaws; 76-27GB, about 0.2 m below the uppermost bed, annelid jaws.

SUTARVE 2. Hemse Beds, lower part or Klinteberg Beds, unit I , top. Age: Early Ludlow. References: Laufeld 1 974a, b' ; Laufeld & Jeppsson 1 976; Jeppsson 1 983; Frykman 1 989. Samples/jawedannelids: 7 1 - 1 02U, ditch exposure, annelid jaws.

SUTARVE 3. Klinteberg Beds, unitl, upper part. Age: Early Ludlow. References: Lauleld 1 974a, bO; Frykman 1 989. Samples/jawed annelids: 75-49GB, the sample belongs somewhere within the uppermost 4 m of unit f, no anne lid jaws.

SVARVARE 1 . Slite Beds, Slite Mari, Pentamerus gothlandicus Beds. Age:Wenlock. References:Laufeld 1 974a, b'; Larsson 1 979; Sivhed 1 976; Bergman 1 980a; Aamskold 1 9S5b. Samples/jawedannelids: 75-6KL, Kettnerites (A.) sisyphivar. val le, annel id jaws; 75-7KL, Kettnerites (K) martinssonii, K (A.) sisyphi var. valle, annelid jaws.

SVARVAA E 3. Slite Beds, Sl ite Mari, Pentamerus gothlandicus Beds. Age:Wenlock. Reference: Larsson 1 979'. Samples/jawed annelids: 8 1 -57U, ditch, Kettnerites (A) sisyphi var. valle, annelid jaws.

SVARVEN 1 . Hegklint Beds unit c. Age: Early Wenlock. References: Laufeld 1 974a, b', Larsson 1 979, Jeppsson 1 983. Sampfes/jawed annelids: 78-3GB, 0.30 m above the reference level, Kettnerites (K) abraham cf. isaac, annelid jaws; 84-34LJ, 2.40-2.20 m below the reference level, Hindenites gladiatus, Keltnerites (K.) martinssonii, K. (K.) abraham isaac, K. (A) sisyphi sisyphi, annelid jaws.

SYSNE 1 . Hemse Beds, unit d(?). Age: Ludlow, Late Leintwardinian.


References: Larsson 1 979'; Jeppsson 1 983; Fredholm 1 988. Samples/jawed annelids: 8 1 -48LJ, at the same level as the road surface, annelid jaws; 8 1 -50LJ , 4 .40 m above 8 1 -48 , fragments.

SYSNEUDD 1 . Hemse Beds, unit d . Age: Ludlow. Reference: Larsson 1 979*. Samples/jawedannelids: 78-5CB, from the base of the bioherm, no annelld jaws; 78-22CB, 0 .30 m above 78-seB, no annelid jaws.

TAll NGS 1 . Slite Beds, unit g . Age: Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979. Samples/jawed anne/ids: 80-2GB, at the same level as the road surface, Kettnerites (K.) cf. martinssonii, K. (A) sisyphi sisyphi, GotJandites slitensis, annelid jaws.

TANGLINGS 2. Hemse Beds, lower-middle part. Age: Ludlow. References: Laufeld 1 974a, b'; Larsson 1 979; Fredholm 1 988. Samples/jawed annelids: 75-89GB, 0.50 m below the surface of the road, annelid jaws; 75-9OC B , 0.20 m above the road surface, annelid jaws.

TANGLINGS KVARN 1 . Hemse Beds, probably middle or upper part. Age: Ludlow. Reference: Stridsberg 1 985*; Fredholm 1 988. Sample/jawed annelids: 83-1 SS, from the upper part of the southern wall of the section, no annelid jaws.

TINGS 1 . Klinteberg Beds, unit f. Age: Early Ludlow. References: Laufeld 1 974a, b*; Larsson 1 979; Frykman 1 989. Samples/jawedannelids: 7 1 -21 4LJ, 0.45-0.55 m below the top of the section, no annelid jaws.

TINGSTÅDE 1. 640453 1 66689 (CK 5760 0 1 84) , ca 5 1 0 m SW of Tingstade church. Topographical map sheet 7 J Fårasund SV & NV. Geological map sheet Aa 1 69 Slite.

Temporary excavation in connection with the construetion of houses, about 20 m NW of the bend in the main road. The sample was collected near the southeasternmost point of the row of small houses. The bluish mari is very rich i n stromatoporoids. Slite Beds, unit gt?). Age: Wenlock. Samp/e/jawed anne/id: 80-6G B , excavated material from about 1 -2 m below ground surface, no annelid jaws.

TJANG OARVE 1 . Hemse Beds, Hemse Mari SE. Age: Ludlow, Leintwardinian. References: Laufeld 1 974a, b*; Larsson 1 979; Fredholm 1 988. Samples/jawed annefids: 7 1 - 1 88LJ, loose slabs on the field, Kettnerites (K.) huberti, annelid jaws.

TJAUTET 1 . Slite Beds, unit e and lower part of unit g . Age: Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979. Sampfes/jawed annelids: Slite Beds, unit e : 72-6LJ, 0.65-0.69 m below the reference level, no annelid jaws; 72-7LJ, 0 .00-0.05 m below the reference level, no annelid jaws.

TJELDERSHOLM 1 . Slite Beds, Pentamerus golhlandicus Beds, and beds immediately younger. Age:Late(?) Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979; Jeppsson 1 983'; Aamskold 1 985b. Samples/jawed annelids: 73-8LJ, 0.35-0.40 m below the reference level , Kettnerites (K) martinssonii, annelid jaws; 73-9U, 0.0-0.07 m below the reference level, Hindenites sp. , Kettnerites (K.) martinssonii, K. (K) jacobi, K. (A) sisyphi sisyphi, Gotlandiles slitensis, annelid jaws; 75-31 GB, about 0.35 m above the reference leve l , Kettnerites (A.) sisyphi sisyphi, Gotlandites s/itensis, annelid jaws; 75-33GB, about 0.35 m above the reference level and 30 m NW of the reference point, Kettnerites (K.) martinssonii, K (A) sisyphi sisyphi, Gotlandites sfitensis, annelid jaws; 75-32GB, 0.40 m above the reference level, GotJancJites slitensis, annelid jaws; 73-1 OU, 0.50-0.55 m above the reference level, Kettnerites (K.) martinssonii, K (A.) sisyphi sisyphi, Gotlandites slitensis, annelid jaws; 73-1 1 LJ , about 3.70 m above the reference level, fragments.

T JULS 3 . Slite Beds, Slite MarI. Age: Wenlock. Reference: Larsson 1 979'. Sampfes/annelid jaws: 84-68LJ, immediately SE of the bridge, ditch exposure, fragments.

TOMASE 1 . Slite Beds, unit g . Age: Wenlock. Reference: Larsson 1 979' ; Ramskold 1 983. Samp/es/jawedannelids: 82-39LJ, south of the road, ditch section, 0.40--0.60 m below the plastic clay layer mentioned by Hede, no annel ids; 82-41 LJ , 0.60-0.70 m above the plastic clay layer, no annefids.

TOMTBODARNE 1. 634368 1 6 6522 (CJ 51 28 41 36), ca 6250 m ESE of Rone church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 56 Ronehamn.

Shallow shore exposure marked on the geological map, about 350 m E of the bend i n the road close to the fishing huts marked on the topographical map sheet. Eke Beds. Age:Ludlow, possibly Late Leintwardinian. Samples/jawed anne/ids: 84- 1 1 OCB, uppermost bed, Kettnerites (K.) martinssonii, K. (K) cf. huberti, K (A) cf. fjaelensis, annelid jaws.

TRADGARDEN 1 . Slite Beds and Halla Beds. Age: Late Wenlock. References: Laufeld 1 974a, b*; Larsson 1 979. Samples/jawed annelids: Slite Beds, Slite Mari, Pentamerus gothlandicus Beds: 7 1 -57LJ, 3.40 m above the base of the sectien and about 1 .60 below the reference level, Kettnerites (K.) polonensis, K. (A) sisyphi sisyphi, annelid jaws.

TRASKE 1 . Hemse Beds, unit b. Age: Ludlow. References: Laufeld 1 974a, b*; Larsson 1 979; Fredholm 1 988.

SampleSljawed annelids: 71-1 03LJ, low ditch exposure, Kettnerites (K.) martinssonii, K. (K) polonensis, K (K) cf. huberti, annelid jaws.

TULE 1 . Klinteberg Beds, unit et?) Age: Latest Wenlock or earliest Ludlow. References: Laufeld 1 974a, b'; Frykman 1 989. Sampfesljawed annelids: 7 1 -95LJ , bed accessible less than 0. 1 -0.2 m below soil surface, paul initid fragments, annelid jaws.

VAKTAAD 2. Hemse Beds, Hemse MarI SE part. Age: Ludlow, Leintwardinian. References: Larsson 1 979'. Samples/jawed annelids: 8 1 -36U, shore exposure, Lanceolatites gracilis, Kettnerites (A.) sisyphi kfasaardensis, K (K) burgensis, annelid jaws.

VAKTARD 3. Hemse Beds, Hemse Mari, SE part. Age: Ludlow, Leintwardinian. Reference: Jeppsson 1 983; Fredholm 1 988. Samples/jawed annelids: 7 1 - 1 49LJ, shallow shore exposure, KetInerites (K.) burgensis, annelid jaws.

VAKTARD 4. 633359 1 646 1 1 (CJ 3 1 68 3275), ca 3 H lQ m WSW of Nås church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 52 Burgsvik.

Ditch exposure south of and along the road, 200 m W of Vaktård 5. Hemse Beds, Hemse Mari S E part. Age: Ludlow, Leintwardinian. Samples/jawedannelids:81 -35LJ, shatlow ditch exposure, Lanceolatites graci/is, Ketlnerites (K.) huberti, K (A) sisyphi k/asaardansis, K. (K) burgensis, annelid jaws.

VAKTARD 5. 633364 1 64651 (CJ 31 68 3276), ca 2400 m WSW of Nås church. Topographical map sheet 5 I Hoburgen NO & 5 J Hemse NV. Geological map sheet Aa 1 52 Burgsvik.

Ditch exposure south of and along the road, 1 50 m W ol the highest point of the road. North of the road is the boundary between pasture and tilled land. Hemse Bads, Hamse Mari SE part. Age: Ludlow, Leintwardinian. Samples/jawedannelids: 8 1 -37LJ , shallow ditch exposure, Lanceofatites gracilis, Kettnerites (A) sisyphi klasaardensis, K. (K) huberti, K (K) burgensis, annelid jaws.

VALBYBODAR 1 . 635895 1 64258 (CJ 2987 5829), ca 1 030 m WSW ol Frejel church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 64 Hemse.

A shore exposure, about 10 m long, just north of the small bay at Valbybodar, 1 75 m NW of the single house marked on the first edition of the topographical map shee1. Slite Beds, Slite Siltstone. Age: Wenlock. Reference: Hede 1 942, Ioc 1 a. Samp/es/jawed annelids: 82-26LJ, 0.2 m below mean water level, Hindenites gladiatus, Kettnerites (K.) cf. bankvaetensis, K (K.) martinssonii, K. (K.) pofonensis, K (A) sisyphicf. var. valle, annelid jaws.

VAL BYTTE 1 . Slite Beds, Slite Mari sl ightly younger than the Rhipidium tenuistriatum Beds. Age: Wenlock. References: Laufeld 1 974a, bO; Larsson 1 979; Ramsk61d 1 983, 1 984; 1 985b. Samples/jawed annelids: 76-45KL, Kettnerites (K.) jacobi. annelid Jaws.

VALBYTTE 2. Slite Beds, Slite Mari sl ightly younger than the Rhipidium tenuistriatum Beds. Age: Wenlock. Reference: Larsson 1 979°. Samples/jawedannelids: 75-30KL, annelid jaws.

VALLE 1 . Slite Beds, Pentamerusgothlandicus Beds. Age:Late(?) Wenlock. References: Laufeld 1 974a, bO; Larssen 1 979. Samples/jawed annelids: 66-1 44SL, 0.0-0 . 1 5 m below top of the section, Kettnerites (A) sisyphivar. valle, annelid jaws.

VALLE 2. Slite Beds, Pantamerusgothlandicus Beds. Age:Late(?) Wenlock. References: Laufeld 1 974a, b* ; Larsson 1 979. SampJes/jawedannelids: 66-1 45SL, 1 .00-1 .25 m below the top of the section, Ketlnerites (K) martinssonii, K (A) microdentatus, K. (A) sisyphivar. valle, annelid jaws.

VALLEV IKEN 1 . Slite Bads, Slite MarI. Age:Wenlock. References: Laufeld 1 974a, b'; Larsson 1 979; Aamskold 1 985b. Samples/jawed annelids: 70- 1 0 LJ . 0.40-0.50 m below the tap of the section i n the northern part, Kettnerites (K) cf. martinssonii, annelid jaws; 70-1 1 LJ, 0. 1 0 m above water level , Kettnerites (A) microdentatus, annelid jaws; 8 1 - 1 GB, sample from the N E part ol the quarry. The bed of which the upper surface is developed as a hardground, covers a large area some metres below the top of the ground surface, Kettnerites (K) bankvaetensis, K (K.) martinssonii, K (A) microdentatus, annelid jaws.

VALLMYR 1 . Klinteberg Beds, unit d . Age:Wenlock, very close to the end. References:Larsson 1 979*; Jeppsson 1 983; Frykman 1 989. Samples/jawed annelids: 75-38LJ, on the northern side of the ditch, Kettnerites (K.) martinssonii, annelid jaws.

VALLSTENA 2. Silte Beds, Pentamerus gothlandicus Beds or sl ightly older. Age: Late(?) Wenlock. References: Laufeld 1 974a, b* ; Larsson 1 979. Samplesljawedannelids: 67-23LJ, excavated material, Kettnerites (K) cf. bankvaetensis, K (K.) cf. martinssonii, Gotlanditesslitensis, annelid jaws; 77-2GB, 0.0-0 . 1 m of the top of the section, Kettnerites (K) cf. bankvaetensis, Gotfandites sfitensis,. Hindenites gladiatus, paul in itid sp. , annelid jaws; 77-3G B, 0.50 m below the top ol


the section, Kettnerites (K.) martinssonii, K. (K.) jacobi, K. (A.) sisyphi sisyphi, Gotlandites slirens/5, Hinden/tes gladiatus, annelid jaws.

VALVE 3. 637322 1 640 1 0 (CJ 2851 7273), ca 2530 m NNW of Vastergarn church. Topographical map sheet 6 I Visby SO. Geological map sheet Aa 1 60 Klintehamn.

Ditch exposure east of the house at the western side of the road running N-S from the cross-roads south of Valve towards Sigvards (NE of Paviken). Slite Bads, unit a. Age:Wenlock. Referenees: Hede 1 927a, p . 24, l ines 34-44. Samplesljawedannelids: 84-66LJ, ditch exposure, Kettnerites (K.) martinssonii, K. (A) microdentatus, K. (A.) sisyphi sisyphi, annelid jaws.

VARSA.NDE 1. Mulda Bads, lowest part. Age: Late Wenlock. References: Laufeld 1 974a, bO. Samples/jawedannelids: 75-23CB, shallow ditch section, Kettnerites (K) martinssonii, K (A) fjaelensis, K (A) sisyphi sisyphi, annelid jaws; 82-41 CB, lowest bed, rich in trilobites, Kettnerites (K.) martinssonii, K (A) microdentatus, annelid jaws; 75-22CB, about 350 m S of the northemmost house at Vårsånde, Kettnerites (K.) martinssonii, K (A) fjaelensis, K (A) sisyphi sisyphi, annelid jaws; 82-42CB, about 400 m S of the northernmost house at Varsande, bottom of the ditch, Kettnerites (K) martinssonii, K. (A) microdentatus, annelid jaws; 82-44CB, loose slab, Mulde Beds, Hindenites sp., Kettnerites (K.) martinssonii var. m ulde, K (A) fjaeJensis, sp. , annelid jaws.

VASTERBACKAR 1 Sundre Beds, middle upper part. Age: Latest Ludlow, Whitcliffian. References: Laufeld 1 974a, bO. Samples/jawed annelids: 7 1 - 1 84LJ, tap of the section, no annelid jaws; 75-2LJ, Kettnerites polonensis, annelid jaws.

VASTERBJARS 1 . Klinteberg Beds. Age: Latest Wenlock - earliest Ludlow. Reference:Frykman 1 989*. Samples/jawedannelids:83-38LJ, from the road-side section north of the cross-roads SW of Vasterbjars. The sample was taken below the road sign 'Horsne', from the lowest accessible bed i n the bottom of the ditch, Kettnerites (K.) bankvaetensis, K. (A) cf. sisyphi, annelid jaws.

VASTLAUS 1 . Hemse Beds, Hemse Mari SE part. Age: Ludlow, probably Leintwardinian. References: Laufeld 1 974a, bO; Larsson 1 979; Fredholm 1 988. Samples/jawed annelids: 82-31 LJ, Lanceolatites gracifis, Kettnerites (K) martinssonii, K. (A) sisyphi klasaardensis, K (K.) huberti, K (K) burgensis, annelid jaws.

VASTOS 1 . Hogklint Beds unit b. Age: Early Wenlock. References: Laufeld 1 974a, bO, Larsson 1 979. Samples/jawedannelids: 77-1 0CB, 3 m below the tap of the surface of the bridge, fragments.

VATE 1 . 637264 1 65427 (CJ 4257 71 07), ca 800 m N E of Vale church. Topographical map sheet 6 J Roma SV. Geological map sheet Aa 1 60 Klinteham n .

Ditch section immediately S E of t h e intersection rcad a n d the ditch of the ca 250 m S E of the cross-roads at Backs. Våte 1 comprises the distance 0-30 m S E of the intersection. Reference point: The small carbonate build-up, about 1 m south of the rcad and below the surface of the road. Halla Beds, unite b(?). Age:Wenlock. SampJes/jawed annelids: 82-35CB, the bed which underlies the carbonate build-up, no anne lid jaws; 82-36CB, about 0.50 m above and 4 m SE of the previous sample and 5 m SE of the road, fragments.

VATE 2. 637373 1 65540 (CJ 4375 7206), ca 2350 m N E of Vale church. Topographical map sheet 6 J Roma SV. Geological map sheet Aa 1 60 Klintehamn .

S u rface exposure 2 m south o f the path and 1 0-15 m west o f a small wooden hut (not marked o n the topographical map). About 900 m N E of the house, at the end of the field road, about 1 .5 km NE of Vate church. Klinteberg Beds, lower part. Age: Late Wenlock. Samplesljawed annelids: 85-40LJ, surface exposure, pau l in itid fragments, annelid fragments.

VATIENFALLSPROFILEN 1 . Lower Visby, Upper Visby, and Hogkl int Beds, units a-d. Age: Early Wenlock. References: Hedstr6m 1 904, p. 93, l ine 1 1 , to p. 96, l ine 1 7, 1 923, p. 1 95, Fig. 2 ; Hede 1 925, p. 1 5, l ine 18 from below, to page 1 6, l ine 3; Martinsson 1 972, pp. 1 28-1 29; Lauleld 1 974b- ; Bassett & Cocks 1 974, p. 5; Laufeld & Jeppsson 1 976; Franzen 1 977, pp. 223, 226; Larsson 1 979; Jaanusson, Laufeld & Skoglund 1 979-; Claesson 1 979; Bengtson 1 981 ; Jeppsson 1 982, 1 983; Brood 1 982; Ramskold 1 983, 1 984, 1 985b. Samples/jawed annelids: Lower Visby Beds unit e : 76-6LJ, 0.94-0.99 m a.s.l., Kettnerites (K) versabilis, Kettnerites (K.) abraham abraham, Lanceolatites gracilis, annelid jaws; Upper Visby Beds: 70-1 4LJ, 0.99-1 .06 m a.s.l . , Kettnerites (K) martinssonii, K. (K.) abraham abraham, annelid jaws; 76-8LJ, 2.58-2.62 m a.s. l . , Lanceolatites gracilis, Kettnerites (Kl abraham isaac, annelid jaws; Hogklint Beds u nit a : 70-20LJ, 1 0 . 02-1 0.04 m a.s . l . , Lanceolatites gracilis, Kettnerites (K.) martinssonii, K (A) microdentatus, K (K) abraham isaac, annelid jaws; Hågklint Beds un it b: 70-8LJ, 1 3.33 m a.s. l . , Lanceolatites gracilis, Kettnerites (K) martinssonii, K (A) microdentatus, annelid jaws; 82-SCB, 1 3.77-1 3.80 m a.s . l . , Kettnerites (K) martinssonii, K (K.) abraham isaac, Lanceolatites gracilis, annelid jaws; 82-6CB, 1 3.80-1 3.83 m a.s. l . , Kettnerites (K) martinssonii, K. (A) microdentatus, K (Kl abraham isaac, Lanceolatites graeilis, annelid jaws; 70-6LJ, 1 5.33-1 5.41 m a.s . l . , Lanceolatites gracilis, Kettnerites (K.) martinssonii, K (A) microdentatus, K. (K) abraham abraham, K. (K.) abraham isaac, Hindenitesgladiatus, annelid jaws; 70-5LJ, 1 6 .0-1 6.1 0 m a.s. l . , Lanæolatites gracilis, Kettnerites (K.) martinssonii, K (A) microdentatus, K. (K.l abraham isaac, annelid jaws; 70-2LJ, 1 9.24-1 9.29 m a.s . l . , Lanceolatites graeilis, Kettnerites (K) martinssonii, K. (A) microdentatus, K (K) abraham isaac, annelid jaws; Hagklint Beds, unit d: light shale RM 29.6-30.0 m a.s.l . , Kettnerites (K) martinssonii, Hindenites angustus, annelid jaws; Valdaria testudo RM 29 .6-30.0 m a.s. l . , Ketlnerffes (K.) martinssonii, K (A) micrcxJentatus, K. (K.) abraham isaac, Hindenites angustus, H. gladiatus; annelid jaws; Herrmannina R M 29.6-30.0 m a.s.l . , Kettnerites (K.) martinssonii, K (A) microdentatus, K. (K) abraham isaac, Hindenites angustus, H. gladiatus, annelid jaws; light limestone RM 29.6-30.0 m a.s. l . , Ketfnerffes (A) sisyphi sisyphi, Hindenltes angustus, annelid jaws; Pterygotus Mari RM 29.6-30.0 m a.s. l . , Kettnerites (K.) martinssonii, K (A) microdentatus, K. (K.) abraham isaac, Hindenites angustus, H. gladiatus, annelid jaws.

VATIENFALLSPROFILEN 2. 639260 1 64823 (CK 3808 9 1 8 1 ) . ca 1 1 00 m SW of Visby cathedral. Topographical map sheet 6 1 Visby NO. Geological map sheet Aa 1 83 Visby & Lummelunda.

Temporary excavation for house construction, about 50 m south of the uppermost part ol the locality Vattenfallsprofilen 1 . The locality is no langer accessible. The exposure comprised many varled lithologies: almost black pyrite-rich calcarenite, red calcarenite, light yellow calcarenite, and reefal l imestone, all found within an excavation roughly 10 m wide and 2-3 m deep. The lithologies formed bodies rather than restricted beds. HogKlint Beds, unit d . Age: Early Wenlock. Samplesljawed annelids: 77-35CB, 77-36CB, and 77-37CB, no annelid jaws; 77-38CB, annelid jaws.

VIALMS 2. 642568 1 69043 (CK 8245 21 1 7) , ca 6820 m N of Bunge church. Topographical map sheet 7 J Får6sund SO & NO. Geological map sheet Aa 1 71 Kappelshamn.


I n /and ditt section W o f Buckhallar and 1 00 m N o f the triangulation point 1 0 ,2 at Vialms. Reference level: The contact between Hogklint Beds and Slite Beds (Hede 1 933, p. 33, l ines 37-40. Hogklint and Slite Beds. Age: Early Wenlock. Reference: Hede 1 933, p. 33, l ine 22, to page 34, l ine 2. Samples/jawed annelids: 84-70LJ, 1 .40-1 .25 below the reference level, fragments.

V I D FALLE 1 . Hemse Beds, unit b . Age: Ludlow. References: Laufeld 1 974a, bO; Larsson 1 979. Samplesljawedannelids: 75-41 CB, 0 . 1 5 m above the bottom of the ditch, Kettnerites (K) martinssonii, K (K) huberti, K. (A) sisyphi, annelid jaws.

VIKE 1 . Slite Beds, Pentamerusgothlandicus or sl ightly older. Age: Late(?) Wenlock. References: Laufeld 1 974a, b-; Larsson 1 979. Samples/jawed annefids: 75-30CB, 0.30 m below the top of the section on the S E side of the bridge, Gotlanditesslitensis, annelid jaws.

VIKE 2. 639466 1 67680 (CJ 6673 91 29), ca 4990 m S of Boge church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 69 Slite.

Drainage ditch section o n the southern side of the ditch , 0-25 m W of the southwestern corner ol the new concrete bridge for road 1 46. Slite Beds, Pentamerus gothlandicus Beds or, more probably, slightly older. Age: Late(?) Wenlock. Reference point: The intersection of the west side of the bridge and the southern side of the ditch. Reference level: The bridge abutment which also forms the bottom of the ditch. Sampleslannelid jaws: 834CB, 0.20-0.40 m above the reference level, Gotlandiles slitensis, Kettnerites (K.) bankvaetensis, K (K) martinssonii, K. (A) microdentatus, Hindenites gladiatus, annelid jaws; 83-5CB, 1 .2-1 .3 m above the reference level , Gotfandites slitensis, annelid jaws.

VIKE 3. 639476 1 67666 (CJ 6660 9 14 1 ), ca 4860 m S of Boge church. Topographical map sheet 6 J Roma NV & NO. Geological map sheet Aa 1 69 Slite.

Drainage ditch lU nning NW-SE, exposure on the southwestern side. Slite Beds, Pentamerus gothlandicus Beds or, more probably, slightly older. Age: Late(?) Wenlock. Reference point: On the southern side of the ditch about 200 m W of the new concrete bridge, a protruding section of about 0.5 cubic metres. Samples/jawed annelids: B3-3CB , about 0.5 m above mean water level, Kettnerites (Kl cf. bankvaetensis, K. (K.) martinssonii, Gotlandites slitensis, Hindenites sp., annelid jaws.

VIVUNGS 1 . Klinteberg Beds, middle-upper part. Age: WenlockLudlow boundary. References: Laufeld 1 974a, b�; Larsson 1 979; Frykman 1 989. Sampfes/jawed annelids: 7B-l 1 CB, 0.2 m Irom the top ol the section, annelid jaws.

YG N E 2. Upper Visby Beds, uppermost part. Age: Early Wenlock. Reference: Laufeld 1 974a, b*; Larsson 1 979. Samples/jawed annelids: 76-1 KL, Kettnerites (K) abraham cf. abraham, annelid jaws.

YXN E 1 . Slite Beds, unit d . Age: Wenlock. Referenees: Lauleld 1 974b* ; Larsson 1 979. Samples/jawed annelids: 76-29CB, excavated material, Kettnerites (K.) cf. bankvaetensis, annelid jaws.

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If there are severai references to any one author, the name should be repeated in each entry. Titles of articles should be decapitalized, except in cases where this would violate a language convention. Cyrillic letters should be transliterated according to the ISO Standard 833 ( 1974) . Serial titles should not be abbreviated.

Figures, plates and table captions should be self-explanatory but as short as possible. lf a figure contains lettered items, list them en suite in the caption and insert a square symbol (may be coded as '$F' ) immediately in front of each entry (OA, etc. ) . All figured specimens should have a reference to their provenance and present location (museum or similar registration number) .

Jllustrations. - All illustrations must be dearly marked with the author's name and figure number. Plan the figures so that they take up the entire width of the type area ( 1 70 mm) or the width of one column (81 mm) . lf an intermediate width has to be used, do not exceed 127 mm. In the event of a full-page illustration, try to allow space for the caption to come within the page depth, 254 mm. Plates should be constructed for an area of l 70x254 mm, but it is usually preferable to arrange all illustrations as figures to be placed in their proper positions in the text.

The cost of reproducing a figure is based on the smallest rectangular frame in which the figure can be inserted. Use that frame! Do not leave open corners or unnecessary space between items, Do not let text of le.tt&ing protrude outside the frame of the figure .

Photographs should be printed on white paper with glossy finish . They must be dear and sharp ly contrasted, but without pronounced light areas and heavy shadows. If incident light is used for iIlumination, the light should fall consistently from the upper left. Stereo-pairs should be mounted at a maximum distance of 70 mm. In a composite figure, all items should be of similar tone and contrast. Composite figures should consist of regular units as far as possible. Mount the photographs on cardboard. If a dean background is desired, provide originals that have an even black background tone or submit overlays for blocking out to an either black or white background (see Bengtson, S. 1986: Preparing dean backgrounds in published photographic iIlustrations. Letkaia 1 9:4, 361-362; available on request from the editor) . If a photographic figure consists of severai rectangular items, make sure that the intervening narrow strips are directly reproducible; i .e . cut the prints accurately and mount them on white cardboard, alternatively edge to edge with white adhesive strips over the joints.

Line drawings should have lines of even thickness and blackening. Do not use gray or toa densely screened (more than 40 lines/ cm) surfaces. If the figures indude text, Do Not Monumentalize the Text by Capitalizing Words. For metric units, use the standard symbols - �, mm, m, km - there should be no capitals (at least not for length units) , not plural , no genitive, no hyphens, and no periods. Separate the symbols from the num ber by a space. Do not leave out zero before decimal points.

Line drawings and half tones should not be combined in the same figure without good reason. Place text and lettering on the figure, normally on its background portions.

Lettering of items in composite figures (A, B, C, etc . ; not a, b, c. .. ) and plates ( 1 , 2 , 3, etc . ) should be distinct but not dominant, and placed as consisten tly as possible in the different items. Use transfer lettering (simple, sans-serif, semibold typefaces such as Helvetica) or stencils.

Fossils and Strata - Foreninger ved UiO

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